CN103080315A - 用于改进植物的农艺学性状的udp-葡萄糖-4-差向异构酶 - Google Patents
用于改进植物的农艺学性状的udp-葡萄糖-4-差向异构酶 Download PDFInfo
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Abstract
公开了UDP-葡萄糖-4-差向异构酶的分离的多核苷酸和多肽和重组DNA构建体,所述UDP-葡萄糖-4-差向异构酶用于改进农艺学性状,包括产量和干旱耐受性。也公开了具有这些重组DNA构建体的转基因植物和使用这些重组DNA构建体的方法。
Description
技术领域
领域涉及作物育种和遗传学,且具体地,涉及可用于在植物中赋予干旱耐受性和产量增加的重组DNA构建体。
背景技术
非生物胁迫是全世界作物损失的主要原因,造成主要作物的超过50%的平均产量损失(Boyer, J.S. (1982) Science 218:443-448; Bray, E.A. 等人(2000) In Biochemistry and Molecular Biology of Plants, Edited by Buchannan, B.B. 等人, Amer. Soc. Plant Biol., 第1158-1249页)。在各种非生物胁迫中,干旱是限制全世界作物产量的一个主要因素。处于不同发育阶段的植物向水限制环境的暴露,似乎会活化多种生理和发育变化。对干旱胁迫感知、转导和耐受性的基础生化和分子机制的理解是生物学的一项重大挑战。
高等植物中的光合反应依赖于叶绿体类囊体膜系统。叶绿体类囊体装配和维护需要膜组分的连续供给。含有半乳糖的甘油脂是植物、藻类和蓝细菌中的光合膜的主要脂质组分。2种最常见的半乳糖脂是单半乳糖基二酰基甘油和二半乳糖基二酰基甘油(MGDG和DGDG),它们分别占总类囊体脂质的约50和25 mol%。所有植物脂质中的约80%与光合膜有关,MGDG被认为是地球上最丰富的膜脂质。半乳糖脂在光合膜的组构中和在它们的光合活性中起重要作用。
在植物中,在2个不同的步骤中合成MGDG:(i) UDP-葡萄糖4-差向异构酶(UGE)将UDP-D-葡萄糖(UDP-Glc)转化成UDP-D-半乳糖(UDP-Gal),和(ii) MGDG合酶(MGD1)将半乳糖基残基从UDP-Gal转移至二酰基甘油(DAG),用于合成终产物。MGD1定位在内叶绿体被膜,并使用UDP-Gal作为底物。
植物具有复杂的糖生物合成机制,所述机制包括这样核苷酸糖家族,所述核苷酸糖可以在它们的糖基部分处被核苷酸糖互变酶修饰以产生不同的糖。UDP-葡萄糖4-差向异构酶(也称作UDP-半乳糖4-差向异构酶, UGE; EC 5.1.3.2)会催化UDP-Glc和UDP-Gal的互变。从植物鉴别出的UGE缺少跨膜基序和信号肽,且似乎作为可溶性的实体存在于细胞质中。通常,植物UDP-Glc差向异构酶定位于胞质溶胶,它们的底物UDP-Glc和UDP-Gal在这里以高水平存在。作为用于在叶绿体中合成半乳糖脂MGDG的前体,UDP-Gal被普遍认为是从胞质溶胶运动,因为UDP-Gal浓度在质体内相对较低,MGDG合酶(MGD1)与内被摸有关。
为了研究控制植物中的光合活性和碳同化的基因,鉴别出了具有减少的光同化物和产量生成的稻生长矮小突变体光同化物缺陷型1 (phd1)。本文公开了一种新颖的定位于叶绿体的UDP-Glc差向异构酶,其涉及光合膜生物发生(biogenesis)过程中叶绿体半乳糖脂生物合成的UDP-Gal供给。
发明内容
公开了质体UDP葡萄糖差向异构酶、它的同系物和使用方法。PHD1的转基因表达会增加光合活性和增强生长。本文讨论了PHD1、其同系物和功能片段在光合能力和碳同化物体内稳态中的作用。
本公开内容包括:
在一个实施方案中,一种植物,所述植物在它的基因组内包含重组DNA构建体,所述构建体包含与至少一个调控元件可操作地连接的多核苷酸,其中所述多核苷酸包含选自下述的核苷酸序列:(a)编码具有质体差向异构酶活性的多肽的核苷酸序列,其中所述多肽具有与SEQ ID NO: 1-17相比至少60%、80%、85%、90%、95%或100%序列同一性的氨基酸序列,这基于Clustal V比对方法,其中逐对比对的默认参数为:KTUPLE=1,GAP PENALTY=3,WINDOW=5和DIAGONALS SAVED=5,(b)编码具有差向异构酶活性的多肽的核苷酸序列,其中所述核苷酸序列可在严谨条件下与包含SEQ ID NO: 18-34的完整互补体的DNA分子杂交,(c)编码具有差向异构酶活性的多肽的核苷酸序列,其中通过至少一种选自缺失、置换、添加和插入的方法,通过改变一个或多个核苷酸,从SEQ ID NO: 18-34衍生出所述核苷酸序列,(d)编码多肽的核苷酸序列,其中所述多肽的氨基酸序列包含SEQ ID NO: 1,和(e)包含SEQ ID NO: 18的核苷酸序列,且其中与不包含所述重组DNA构建体的对照植物相比,所述植物表现出增加的干旱耐受性。所述植物可以是单子叶植物或双子叶植物。
在一个实施方案中,所述PHD1多肽不具有SEQ ID NO: 1的N-端叶绿体转运肽1-62氨基酸或在其它PHD1同系物中的对应等效肽。例如,在植物细胞中(例如,在质体中),表达基本上缺少编码叶绿体转运肽的区域的核苷酸分子。
在一个实施方案中,将叶绿体转运肽(SEQ ID NO: 1的1-62氨基酸,或与SEQ ID NO: 1的62个氨基酸的N-端区域基本上类似的序列)与用于将表达的蛋白/肽运输进叶绿体中的异源肽融合。
在另一个实施方案中,一种植物,所述植物在它的基因组内包含重组DNA构建体,所述构建体包含与至少一个调控元件可操作地连接的多核苷酸,其中所述多核苷酸包含选自下述的核苷酸序列:(a)编码具有差向异构酶活性的多肽的核苷酸序列,其中所述多肽具有与SEQ ID NO: 1-17相比至少60%、80%、85%、90%、95%或100%序列同一性的氨基酸序列,这基于Clustal V比对方法,其中逐对比对的默认参数为:KTUPLE=1,GAP PENALTY=3,WINDOW=5和DIAGONALS SAVED=5,(b)编码具有差向异构酶活性的多肽的核苷酸序列,其中所述核苷酸序列可在严谨条件下与包含SEQ ID NO: 18-34的完整互补体的DNA分子杂交,(c)编码具有差向异构酶活性的多肽的核苷酸序列,其中通过至少一种选自缺失、置换、添加和插入的方法,通过改变一个或多个核苷酸,从SEQ ID NO: 18-34衍生出所述核苷酸序列,(d)编码多肽的核苷酸序列,其中所述多肽的氨基酸序列包含SEQ ID NO: 1,和(e)包含SEQ ID NO: 18的核苷酸序列,且其中与不包含所述重组DNA构建体的对照植物相比,所述植物表现出产量增加。在水限制条件下,与不包含所述重组DNA构建体的对照植物相比,所述植物可以表现出所述产量增加。所述植物可以是单子叶植物或双子叶植物。
在另一个实施方案中,一种增加植物的干旱耐受性的方法,所述方法包括:(a)向可再生的植物细胞中导入重组DNA构建体,所述构建体包含与至少一个调控元件可操作地连接的多核苷酸,其中所述多核苷酸选自下述的核苷酸序列:(i)编码具有差向异构酶活性的多肽的核苷酸序列,其中所述多肽具有与SEQ ID NO: 1-17相比至少60%、80%、85%、90%、95%或100%序列同一性的氨基酸序列,这基于Clustal V比对方法,其中逐对比对的默认参数为:KTUPLE=1,GAP PENALTY=3,WINDOW=5和DIAGONALS SAVED=5,(ii)编码具有差向异构酶活性的多肽的核苷酸序列,其中所述核苷酸序列可在严谨条件下与包含SEQ ID NO: 18-34的完整互补体的DNA分子杂交,(iii)编码具有差向异构酶活性的多肽的核苷酸序列,其中通过至少一种选自缺失、置换、添加和插入的方法,通过改变一个或多个核苷酸,从SEQ ID NO: 18-34衍生出所述核苷酸序列,(iv)编码多肽的核苷酸序列,其中所述多肽的氨基酸序列包含SEQ ID NO: 1,和(v)包含SEQ ID NO: 18的核苷酸序列;和(b)从步骤(a)以后的可再生的植物细胞再生转基因植物,其中所述转基因植物在它的基因组中包含所述重组DNA构建体,并表现出与不包含所述重组DNA构建体的对照植物相比增加的干旱耐受性。所述方法可以另外包括:(c)获得从所述转基因植物衍生出的后代植物,其中所述后代植物在它的基因组中包含所述重组DNA构建体,并表现出与不包含所述重组DNA构建体的对照植物相比增加的干旱耐受性。
在另一个实施方案中,一种评价植物的干旱耐受性的方法,所述方法包括:(a)获得转基因植物,其中所述转基因植物在它的基因组中包含重组DNA构建体,所述构建体包含与至少一个调控元件可操作地连接的多核苷酸,其中所述多核苷酸包含选自下述的核苷酸序列:(i)编码具有差向异构酶活性的多肽的核苷酸序列,其中所述多肽具有与SEQ ID NO: 1-17相比至少60%、80%、85%、90%、95%或100%序列同一性的氨基酸序列,这基于Clustal V比对方法,其中逐对比对的默认参数为:KTUPLE=1,GAP PENALTY=3,WINDOW=5和DIAGONALS SAVED=5,(ii)编码具有差向异构酶活性的多肽的核苷酸序列,其中所述核苷酸序列可在严谨条件下与包含SEQ ID NO: 18-34的完整互补体的DNA分子杂交,(iii)编码具有差向异构酶活性的多肽的核苷酸序列,其中通过至少一种选自缺失、置换、添加和插入的方法,通过改变一个或多个核苷酸,从SEQ ID NO: 18-34衍生出所述核苷酸序列,(iv)编码多肽的核苷酸序列,其中所述多肽的氨基酸序列包含SEQ ID NO: 1,和(v)包含SEQ ID NO: 18的核苷酸序列和(b)获得从(a)的转基因植物衍生出的后代植物,其中所述后代植物在它的基因组中包含所述重组DNA构建体;和(c)与不包含所述重组DNA构建体的对照植物相比,评价所述后代植物的干旱耐受性。
在另一个实施方案中,一种测定植物的农艺学特性的改变的方法,所述方法包括:(a)获得转基因植物,其中所述转基因植物在它的基因组中包含重组DNA构建体,所述构建体包含与至少一个调控元件可操作地连接的多核苷酸,其中所述多核苷酸选自下述的核苷酸序列:(i)编码具有差向异构酶活性的多肽的核苷酸序列,其中所述多肽具有与SEQ ID NO: 1-17相比至少60%、80%、85%、90%、95%或100%序列同一性的氨基酸序列,这基于Clustal V比对方法,其中逐对比对的默认参数为:KTUPLE=1,GAP PENALTY=3,WINDOW=5和DIAGONALS SAVED=5,(ii)编码具有差向异构酶活性的多肽的核苷酸序列,其中所述核苷酸序列可在严谨条件下与包含SEQ ID NO: 18-34的完整互补体的DNA分子杂交,(iii)编码具有差向异构酶活性的多肽的核苷酸序列,其中通过至少一种选自缺失、置换、添加和插入的方法,通过改变一个或多个核苷酸,从SEQ ID NO: 18-34衍生出所述核苷酸序列,(iv)编码多肽的核苷酸序列,其中所述多肽的氨基酸序列包含SEQ ID NO: 1,和(v)包含SEQ ID NO: 18的核苷酸序列和(b)获得从步骤(a)的转基因植物衍生出的后代植物,其中所述后代植物在它的基因组中包含所述重组DNA构建体;和(c)测定所述后代植物与不包含所述重组DNA构建体的对照植物相比是否表现出至少一种农艺学特性的改变。所述测定步骤(c)可以包括:在水限制条件下,与不包含所述重组DNA构建体的对照植物相比,测定所述转基因植物是否表现出至少一种农艺学特性的改变。所述至少一种农艺学特性可以是产量,而且可以是产量增加。
在另一个实施方案中,一种分离的多核苷酸,其包含选自下述的核苷酸序列:(a)编码具有差向异构酶活性的多肽的核苷酸序列,其中所述多肽具有与SEQ ID NO: 1-17相比至少60%、80%、85%、90%或95%序列同一性的氨基酸序列,这基于Clustal V比对方法,其中逐对比对的默认参数为:KTUPLE=1,GAP PENALTY=3,WINDOW=5和DIAGONALS SAVED=5,(b)编码具有差向异构酶活性的多肽的核苷酸序列,其中所述核苷酸序列可在严谨条件下与包含SEQ ID NO: 18-34的完整互补体的DNA分子杂交,(c)编码具有差向异构酶活性的多肽的核苷酸序列,其中通过至少一种选自缺失、置换、添加和插入的方法,通过改变一个或多个核苷酸,从SEQ ID NO: 18-34衍生出所述核苷酸序列,(d)编码多肽的核苷酸序列,其中所述多肽的氨基酸序列包含SEQ ID NO: 1,和(e)包含SEQ ID NO: 18的核苷酸序列。
在另一个实施方案中,一种分离的多核苷酸,其包含本公开内容的核苷酸序列的完整互补体,其中所述完整互补体和本公开内容的核苷酸序列由相同数目的核苷酸组成,且是100%互补的。
在另一个实施方案中,一种重组DNA构建体,其包含与至少一个调控元件可操作地连接的、本公开内容的分离的多核苷酸。
在另一个实施方案中,一种细胞,其包含本公开内容的重组DNA构建体,其中所述细胞选自:细菌细胞、酵母细胞和昆虫细胞和植物细胞。
在另一个实施方案中,一种植物或种子,其包含本公开内容的重组DNA构建体。所述植物或种子可以是单子叶植物或双子叶植物的植物或种子。
在另一个实施方案中,一种用于分离由本公开内容的重组DNA构建体编码的多肽的方法,其中所述方法包括下述步骤:(a)用本公开内容的重组DNA构建体转化细胞;(b) 在适合表达所述重组DNA构建体的条件下,培养步骤(a)的转化的细胞;和(c)从步骤(b)的转化的细胞中分离所述多肽。
在另一个实施方案中,一种载体,其包含本公开内容的多核苷酸。
在另一个实施方案中,一种用于生产转基因植物的方法,所述方法包括:用本公开内容的重组DNA构建体转化植物细胞,和从所述转化的植物细胞再生转基因植物。
在另一个实施方案中,本公开内容包括本公开内容的任一种植物,其中所述植物选自:玉米、大豆、向日葵、高粱、油菜、小麦、苜蓿、棉花、稻、大麦、粟、甘蔗、柳枝稷、烟草、马铃薯和糖用甜菜。
在另一个实施方案中,本公开内容包括本公开内容的任一种方法,其中所述植物选自:玉米、大豆、向日葵、高粱、油菜、小麦、苜蓿、棉花、稻、大麦、粟、甘蔗、柳枝稷、烟草、马铃薯和糖用甜菜。
在另一个实施方案中,本公开内容包括本公开内容的任一种植物的种子,其中所述种子在它的基因组中包含重组DNA构建体,所述构建体包含与至少一个调控元件可操作地连接的多核苷酸,其中所述多核苷酸编码多肽,所述多肽具有与SEQ ID NO: 1相比至少60%序列同一性的氨基酸序列,这基于Clustal V比对方法,且其中从所述种子生产的植物表现出与不包含所述重组DNA构建体的对照植物相比增加的干旱耐受性或产量增加或二者。
附图和序列表说明
从下面的详细描述以及附图和序列表中,可以更完整地理解本公开内容,所述附图和序列表形成本申请的一部分。
图1显示了phd1突变体表型。(A)在MS培养基上生长的2周龄幼苗。(B和C)在水稻田中生长的2月龄(B)和4月龄(C)植物的生长表型。(D)收获的圆锥花序表现出降低的phd1-1结籽率。(A-D)野生型(左)和phd1-1 (右)。(E-G)每株植物分蘖数(E)、结实率(F)和粒重(G)的农业特性定量。每个条是30个重复样品的平均值±标准差。(H, I) phd1-1和野生型的蔗糖(H)和淀粉(I)含量的昼夜变化。收获处于开花期的各个野生型和phd1-1植物的成熟叶子,并立即在液氮中冷冻。每个点是10个重复样品的平均值±标准差。
图2显示了PHD1的分子鉴别。(A) PHD1基因的结构和它在3个phd1等位基因中的突变位点。PHD1基因由9个外显子(绿框)和8个内含子(灰线)组成。指示了在3个phd1等位基因中的核苷酸插入和置换。(B, C) phd1突变体的功能互补。(B)处于分蘖期的野生型、phd1-1和互补系phd1-1+PHD1植物的表型。通过半定量RT-PCR,检测PHD1转录物的表达水平。(C)处于开花期的野生型、phd1-1和互补系phd1-1+PHD1植物在白天正午时旗叶中的蔗糖和淀粉含量。误差棒代表8个不同个体的标准差。* phd1-1突变体和野生型之间的显著差异(P=0.05)。
图3显示了PHD1的系统进化分析。从绿色植物中的PHD1 (Os01g0367100)同系物的氨基酸序列,推断出MEGA4邻近连接树。自展值(Bootstrap value)是基于1000次复制,并指示在它们各自的结点处。比例条指示基于分支长度的遗传距离。构建的树的比对显示在图S3中。
图4显示了PHD1表达分析和PHD1蛋白亚细胞定位。(A)在即将到开花期之前,根、秆、花、叶片和叶鞘中的PHD1基因的RNA凝胶印迹分析。(B-E)通过mRNA原位杂交检测的PHD1表达谱。在茎干顶端分生组织和嫩叶(B)、维管束周围的叶肉细胞(C)、幼花序(D)和腋芽(E)处检测到的PHD1信号。在(B)、(C)和(E)中,条= 150ìm在(D)中条= 500ìm。(F)用35S::GFP (下图)和35S::PHD1-GFP (上图)质粒转化的稻原生质体。叶绿素自发荧光(中);GFP荧光(左);合并的图像(右)。条= 5ìm。
图5显示了在phd1-1植物的叶子的UDP-Gal生物合成和碳水化合物分配中涉及的关键基因的表达分析。(A)在UDP-Gal生物合成途径中涉及的基因的表达在phd1-1中被增量调节。(B)淀粉生物合成基因的表达在phd1-1中被减量调节。(C)蔗糖裂解基因的表达在phd1-1中被增量调节。将表达值显示为相对于稻18S RNA的表达比(平均值±标准误)。用3个生物学重复样本进行所有试验。
图6显示了phd1植物中的UDP-糖含量和糖脂组成。(A)在野生型、phd1-1和PHD1-补足的植物的叶子中的UDP-Glc和UDP-Gal含量。所述值代表6个独立实验的平均值±标准误。(B)在水稻田中生长的野生型、phd1-1和PHD1-补足的植物的叶子中的极性脂质组成。通过TLC分离叶脂质,通过GC定量甘油脂。所述条显示了以mol%计的脂质组成,并指示3次测量的平均值±标准差。UDP-Glc,UDP-葡萄糖;UDP-Gal,UDP-半乳糖;FW,鲜重;PC,磷脂酰胆碱;PE,磷脂酰乙醇胺;PG,磷脂酰甘油;PI,磷脂酰肌醇;SQDG,磺基奎诺糖基二酰基甘油。
图7显示了过表达PHD1的转基因稻的改良的农业特性。(A)在水稻田中生长的处于灌浆期的野生型、phd1-1和转基因系S3的表型差异。(B)野生型、phd1-1和过表达PHD1的转基因系(S3、S5和S8)中的PHD1蛋白表达的免疫印迹分析。显示了微管蛋白作为加载对照。在转基因系中观察到增加的PHD1蛋白积累。(C,D)在转基因植物中,每株籽粒产量(C)和收获后营养器官干重(D)增加。值是得自至少30个植物/系的平均值±标准差。* 显著差异(P < 0.05)。
图8显示了PHD1在叶绿体膜的半乳糖脂生物合成途径中的作用的示意模型。与包含PHD1的途径一起,指示了在正常生长条件下的2种甘油糖脂的生物合成路线图。Glc-1-P,葡萄糖-1-磷酸;UGP,UDP-葡萄糖焦磷酸化酶;DAG,二酰基甘油;UTP,尿苷-5’-三磷酸盐;PPi,焦磷酸盐。
图9显示了phd1-1和WT的己糖浓度的昼夜变化。收获各个野生型(●黑色符号+实线)和phd1-1 (○空心符号+虚线)植物的成熟叶子,并立即在液氮中冷冻。每个点是10个重复样品的平均值±标准差。
图10显示了在野生型和3个等位基因phd1突变体中的PHD1转录物水平。通过ACTIN1转录物的RT-PCR检测,证实了样品中RNA的相同丰度。phd1-1至-3,3个等位基因phd1突变体系;S3-1,过表达PHD1的转基因系。
图11是得自PHD1和绿色植物同系物序列的17个保守区的对比。使用BLASTP,搜索所述序列,并使用CLUSTALW进行比对。相同的氨基酸残基带有框,类似的残基带有阴影。红色条指示保守基序GXGXXG (NAD+-结合),活性部位的催化氨基酸残基带有红色框。PHD1: Os01g0367100。
图12显示了PHD1的生化功能和遗传互补试验。(A) PHD1的体外UGE活性试验。纯化的重组PHD1 UGE活性在30℃(■)和在37℃(□)时的利-伯二氏标绘图。(B) PHD1可以互补酿酒酵母gal10突变体。用含有PHD1 cDNA的质粒转化酵母gal10突变株,并在葡萄糖或半乳糖培养基上培养。
序列描述和所附序列表符合在37 C.F.R. §1.821-1.825中阐述的关于专利申请的核苷酸和/或氨基酸序列公开的规则。
序列表含有核苷酸序列字符的单字母代码和氨基酸的3字母代码,它们的定义与通过引用并入本文中的Nucleic Acids Res. 13:3021-3030 (1985)和Biochemical J. 219 (No. 2):345-373 (1984)所述的IUPAC-IUBMB标准相一致。为核苷酸和氨基酸序列数据使用的标识和格式符合在37 C.F.R. §1.822中所述的规则。
具体实施方式
PHD1编码参与叶绿体半乳糖脂的生物合成的新颖的稻质体UGE。在PHD1中的突变会导致被扰乱的碳同化体内稳态和受损的光合效率。PHD1编码定位于叶绿体的有活性的差向异构酶,因此,可以在叶绿体中在原位产生用于MGDG生物合成的UDP-Gal底物(图8)。该令人惊讶的结果提供了一种遗传和生化框架,以工程化质体中的该UGE的新颖功能机制,并评价半乳糖脂在植物(包括稻)的光合活性中的作用。
MGD1被视作对于类囊体膜生物发生而言最重要的MGDG合酶的一种同种型。在拟南芥属中,另外两种MGDG合酶MGD2和MGD3被靶向外叶绿体被膜,在此处可以从胞质溶胶补充底物。由它们产生的MGDG可以从外被膜移动至内被膜和移动至类囊体。在本文中,经证实,与野生型相比,在phd1-1中的主要半乳糖脂MGDG的相对量减少了19%,而DGDG的相对量仅轻微减少了2.5%。但是,发现几种磷脂的mol% 量的轻微增加会补偿在phd1-1突变体中丧失的大约7 mol%的半乳糖脂。这些结果与下述观念相一致:在植物中相反地控制糖脂和磷脂的量,以维持类囊体膜中的脂质的适当平衡。
在正常生长和发育过程中,大部分半乳糖脂定位于质体膜,但是,在磷酸盐(Pi)饥饿以后,DGDG还可以存在于质体外膜中。另外,在蓝细菌中的光合蛋白的x-射线结晶分析,揭示了MGDG与光系统I和II(PSI和PSII)的反应中心的核心有关,这表明,这些脂质不仅作为光合膜的主要组分而被需求,而且被用于光合反应本身。在phd1-1突变体中,光系统II的光化学能量转换的有效量子产率(?PSII)下降。缺少MGDG的幼苗被证实具有破裂的光合膜,这会导致光合能力和光合自养生长的完全损害。MGDG在phd1-1突变体中下降至野生型水平的80%,会导致生长缓慢、光合能力下降和光同化物积累减少的显著表型。phd1-1突变体的生长矮小表型可以归因于,在植物生长过程中支持叶绿体增殖的膜结构单元的供给不足,这也与突变体植物中的叶绿体尺寸减小相一致。这些效应可以归因于叶绿体膜中的MGDG的绝对量的减少或半乳糖脂与磷脂之比的减小。
在植物中,淀粉的作用是,作为白天在叶子中生产的还原碳的仓库,并作为夜间化学能和合成代谢源分子的补给。在phd1-1突变体中,淀粉生物合成基因(诸如AGP、SS、GBSS和BE)在源叶中的表达水平显著地减量调节,这会导致淀粉含量的急剧下降。但是,减少的淀粉不会导致增加的蔗糖水平,因为蔗糖裂解基因SuSy1和INV1/3的活化会导致减少的蔗糖和增加的己糖-磷酸和UDP-Glc水平。因此,蔗糖作为在源器官中生产的光同化物的主要运输形式,不能有效地运输至库器官。此外,由SuSy1或UGP2催化的大量UDP-Glc会被细胞溶质的OsUGE1/4转化成UDP-Gal,并作为叶绿体糖脂的半乳糖基供体运输进叶绿体中,以补偿phd1-1突变体中的PHD1活性损失。相比而言,在稻中的PHD1过表达(其会增强叶绿体中的PHD1活性,且可能增加MGDG的相对量,并增加类囊体膜中的光化学能量转换的有效量子产率)会导致增加的光合效率和生长速率,这暗示PHD1在植物(包括稻)的光合系统中的关键作用。生物量生产和籽粒产量的这些提高,在传统的作物改良和生物能作物生产中都具有重大的经济意义。
本文所述的每篇参考文献的公开内容通过引用整体并入本文。
如在本说明书和所附权利要求书中所使用的,单数形式“一种”、“一个”和“该”包括复数对象,除非上下文清楚地作出相反说明。因而,例如,“一种植物”包括多种这样的植物,“一个细胞”包括1个或多个细胞和本领域技术人员已知的其等效物,依此类推。
如本文使用的:
多肽的“差向异构酶活性”是指,能够执行UDP-Gal向UDP-Glc的催化的肽。
术语“单子叶植物”和“单子叶的植物”在本文中可互换地使用。本公开内容的单子叶植物包括禾本科。
术语“双子叶植物”和“双子叶的植物”在本文中可互换地使用。本公开内容的双子叶植物包括下述科:十字花科、豆科和茄科。
术语“完整互补体”和“全长互补体”在本文中可互换地使用,并表示给定的核苷酸序列的互补体,其中所述补体和所述核苷酸序列由相同数目的核苷酸组成,且是100%互补的。
“拟南芥属”和“拟南芥”在本文中可互换地使用,除非另外指出。
“已表达序列标志”(“EST”)是从cDNA文库衍生出的DNA序列,并因此是已经转录的序列。EST通常通过cDNA插入片段的单个测序通路而得到。整个cDNA插入片段的序列称作“全插入物序列”(“FIS”)。“毗连群”序列是从2个或更多个序列(其可以选自、但不限于EST、FIS和PCR序列)装配成的序列。编码整个蛋白或功能蛋白的序列被称作“完整基因序列”(“CGS”),且可以从FIS或毗连群衍生出。
“农艺学特性”是一种可测量的参数,包括、但不限于:绿度(greenness)、产量、生长速率、生物量、成熟鲜重、成熟干重、果实产量、种子产量、总植物氮含量、果实氮含量、种子氮含量、营养组织中的氮含量、总植物游离氨基酸含量、果实游离氨基酸含量、种子游离氨基酸含量、营养组织中的游离氨基酸含量、总植物蛋白含量、果实蛋白含量、种子蛋白含量、营养组织中的蛋白含量、干旱耐受性、氮吸收、根倒伏、收获指数、茎倒伏、植物高度、穗高度、穗长度、盐耐受性、早期幼苗活力和在低温胁迫下的出苗。
“转基因的”是指,其基因组已经因为存在异源核酸而改变的任意细胞、细胞系、愈伤组织、组织、植物部分或植物,所述异源核酸是例如重组DNA构建体,包括那些最初的转基因事件以及从最初的转基因事件通过有性杂交或无性繁殖建立的那些。本文使用的术语“转基因的”不包括通过常规植物育种方法或通过天然存在的事件对基因组(染色体的或染色体外)的改变,所述天然存在的事件例如随机的异体受精、非重组病毒感染、非重组细菌转化、非重组转座或自发突变。
当应用于植物细胞时,“基因组”不仅包括在核内发现的染色体DNA,而且包括在细胞的亚细胞组分(例如,线粒体、质体)内发现的细胞器DNA。
“植物”包括完整植物、植物器官、植物组织、种子和植物细胞和它们的后代。植物细胞包括、但不限于,来自种子、悬浮培养物、胚、分生组织区域、愈伤组织、叶、根、芽、配子体、孢子体、花粉和小孢子的细胞。
“后代”包含植物的任意后续世代。
“转基因植物”包括在基因组内包含异源多核苷酸的植物。例如,所述异源多核苷酸稳定整合入基因组中,使得该多核苷酸传递连续世代。所述异源多核苷酸可以单独地或作为重组DNA构建体的一部分整合入基因组中。
“异源”序列是指源自外来物种的序列,或者,如果来自相同物种,通过故意的人工干预组成和/或基因组基因座从它的天然形式显著修饰。
“多核苷酸”、“核酸序列”、“核苷酸序列”或“核酸片段”可互换使用,是单链或双链RNA或DNA的聚合物,其任选地含有合成的、非天然的或改变的核苷酸碱基。核苷酸(通常以它们的5'-单磷酸形式存在)用如下的单字母命名来表示:“A”表示腺苷酸或脱氧腺苷酸(分别对于RNA或DNA),“C”表示胞苷酸或脱氧胞苷酸,“G”表示鸟苷酸或脱氧鸟苷酸,“U”表示尿苷酸,“T”表示脱氧胸苷酸,“R”表示嘌呤(A或G),“Y”表示嘧啶(C或T),“K”表示G或T,“H”表示A或C或T,“I”表示肌苷,且“N”表示任意核苷酸。
“多肽”、”肽”、“氨基酸序列”和”蛋白”在本文中可互换使用,是指氨基酸残基的聚合物。该术语应用于氨基酸聚合物,其中的一个或多个氨基酸残基是对应的天然存在的氨基酸的人工化学类似物,也应用于天然存在的氨基酸聚合物。术语”多肽”、”肽”、“氨基酸序列”和”蛋白”也包括修饰,包括、但不限于,糖基化、脂质结合、硫酸化、谷氨酸残基的γ-羧基化、羟基化和ADP-核糖基化。
“信使RNA (mRNA)”指没有内含子、且可以被细胞翻译成蛋白的RNA。
“cDNA”是指与mRNA模板互补且使用反转录酶从mRNA模板合成的DNA。cDNA可以是单链的,或可以使用DNA聚合酶I的克列诺片段转化成双链形式。
“成熟的”蛋白指翻译后加工的多肽;即,已经从其中去除最初翻译产物中存在的任意前肽或肽原的多肽。
“前体”蛋白指mRNA翻译的最初产物;即,前肽和肽原仍然存在。前肽和肽原可以是,但不限于,细胞内定位信号。
“分离的”指以下材料,例如核酸分子和/或蛋白,其基本上不含有或去除了在天然存在的环境中通常伴随该材料或与该材料相互作用的组分。分离的多核苷酸可以从它们天然存在的宿主细胞纯化而来。可以使用技术人员已知的常规核酸纯化方法,来获得分离的多核苷酸。该术语也包括重组多核苷酸和化学合成的多核苷酸。
“重组”指2个原本是分开的序列区段的人工组合,例如,通过化学合成或通过遗传工程技术操纵分离的核酸区段。”重组”也包括已经通过导入异源核酸而修饰的细胞或载体,或源自这样修饰的细胞的细胞,但是不包括通过天然存在的事件(例如,自发突变、天然转化/转导/转座)对细胞或载体的改变,例如不经故意的人工干预而产生的那些。
“重组DNA构建体”是指天然通常不会一起发现的核酸片段的组合。因此,重组DNA构建体可以包含源自不同来源的调节序列和编码序列,或源自相同来源的调节序列和编码序列,但是以不同于天然通常发现的方式排列。
术语“输入克隆”和“输入载体”在本文中可互换地使用。
“调节序列”是指位于编码序列的上游(5'非编码序列)、内部或下游(3'非编码序列)的核苷酸序列,其会影响相关的编码序列的转录、RNA加工或稳定性或翻译。调节序列可以包括、但不限于,启动子、翻译先导序列、内含子和聚腺苷酸化识别序列。术语“调节序列”和“调控元件”在本文中可互换地使用。
“启动子”是指能控制另一个核酸片段的转录的核酸片段。
“在植物中有功能的启动子”是能控制植物细胞中的转录的启动子,无论它的起源是否来自植物细胞。
“组织特异性的启动子”和“组织偏爱型启动子”可互换地使用,是指在一种组织或器官中优势地、但是不一定排它地表达的启动子,但是其也可能在一种特定细胞中表达。
“发育调节的启动子”是指,其活性由发育事件决定的启动子。
“可操作地连接的”指单个片段中核酸片段的缔合,使得一个片段的功能受到其它片段的调节。例如,当能调节核酸片段的转录时,启动子与该核酸片段可操作地连接。
“表达”指功能产物的生成。例如,核酸片段的表达可以指核酸片段的转录(例如,产生mRNA或功能性RNA的转录)和/或mRNA翻译为前体或成熟蛋白。
“表型”是指细胞或生物体的可检测的特征。
在将核酸片段(例如,重组DNA构建体)插入细胞中的上下文中,“导入”指“转染”或“转化”或“转导”,包括核酸片段向真核或原核细胞中的掺入,其中所述核酸片段可以掺入细胞的基因组(例如,染色体、质粒、质体或线粒体的DNA)中,转化成自主复制子,或瞬时表达(例如,转染的mRNA)。
“转化的细胞”是其中已经导入了核酸片段(例如,重组DNA构建体)的任意细胞。
本文使用的“转化”指稳定转化和瞬时转化。
“稳定转化”指核酸片段向宿主生物体基因组中的导入,导致遗传上稳定的遗传。一旦稳定转化,将核酸片段稳定整合入宿主生物体和任意后代的基因组中。
“瞬时转化”是指核酸片段向宿主生物体的细胞核、或含有DNA的细胞器中的导入,导致没有遗传上稳定的遗传的基因表达。
“等位基因”是占据染色体上的特定基因座的基因的几种替代形式之一。当在二倍体植物的一对同源染色体上的特定基因座处存在的等位基因相同时,该植物在该基因座处是纯合的。如果在二倍体植物的一对同源染色体上的特定基因座处存在的等位基因不同,该植物在该基因座处是杂合的。如果转基因存在于二倍体植物的一对同源染色体之一上,该植物在该基因座处是半合的。
“叶绿体转运肽”是这样的氨基酸序列:其与蛋白一起翻译,并将所述蛋白引导至在特定细胞(在其中生产所述蛋白)内存在的叶绿体或其它质体类型。“叶绿体转运序列”是指,编码叶绿体转运肽的核苷酸序列。“信号肽”是这样的氨基酸序列:其与蛋白一起翻译,并将所述蛋白引导至分泌系统(Chrispeels (1991) Ann. Rev. Plant Phys. Plant Mol. Biol. 42:21-53)。如果所述蛋白要导向液泡,可以另外添加液泡靶向信号(出处同上),或者,如果所述蛋白要导向内质网,可以添加内质网保留信号(出处同上)。如果所述蛋白要导向细胞核,将除去存在的任意信号肽,并包括替代的细胞核定位信号(Raikhel (1992) Plant Phys. 100:1627-1632)。“线粒体信号肽”是将前体蛋白引导进线粒体中的氨基酸序列(Zhang和Glaser (2002) Trends Plant Sci 7:14-21)。
通过目检和数学计算,可以确定2个氨基酸或核酸序列之间的同一性百分比。
或者,使用许多用于检测同源序列的对比方法,包括、但不限于,LASERGENE?生物信息计算包(DNASTAR?Inc., Madison, WI)的MEGALIGN?程序,可以测定序列比对和同一性百分比计算。除非另有说明,使用Clustal V比对方法(Higgins和Sharp (1989) CABIOS. 5:151 153),利用默认参数(GAP PENALTY=10, GAP LENGTH PENALTY=10),进行本文提供的序列的多个比对。使用Clustal V方法逐对比对和计算蛋白序列的同一性百分比的默认参数是:KTUPLE=1,GAP PENALTY=3,WINDOW=5和DIAGONALS SAVED=5。对于核酸,这些参数是KTUPLE=2,GAP PENALTY=5,WINDOW=4和DIAGONALS SAVED=4。比对序列后,通过在相同程序上观察“序列距离”表,使用Clustal V程序,可以得到“同一性百分比”和“趋异性”值;除非另有说明,以该方式计算在本文中提供和要求保护的同一性百分比和趋异性。
或者,基于Needleman, S. B.和Wunsch, C. D.的算法(J. Mol. Biol., 48:443-453, 1970),和使用可从威斯康辛大学遗传学计算机组(University of Wisconsin Genetics Computer Group,UWGCG)得到的GAP计算机程序,通过对比序列信息,可以测定2个蛋白序列之间的同一性百分比。GAP程序的优选默认参数包括:(1)评分矩阵,blosum62,参见Henikoff, S.和Henikoff, J. G. (Proc. Natl. Acad. Sci. USA, 89:10915-10919, 1992);(2) 12的间隙权重;(3) 4的间隙长度权重;和(4)对末端间隙没有罚分。
还可以使用本领域技术人员用于序列对比的其它程序。例如,通过使用Altschul等人(Nucl. Acids. Res., 25, 第3389-3402页, 1997)描述的BLAST程序,通过对比序列信息,可以测定同一性百分比。该程序可在因特网上在国家生物技术信息中心(National Center for Biotechnology Information, NCBI)或日本DNA数据库(DNA Data Bank of Japan,DDBJ)的网站得到。在这些网站上,显示了使用BLAST程序进行同一性搜索时的各种条件(参数)的细节,默认值常用于搜索,尽管在适当时可以改变部分设置。或者,使用诸如遗传信息处理软件GENETYX Ver.7 (Genetyx Corporation, Japan)等程序,或使用诸如FASTA等算法,可以测定2个氨基酸序列之间的同一性百分比。在该情况下,默认值可以用于搜索。
通过目检和数学计算,可以测定2个核酸序列之间的同一性百分比,或更优选地,通过使用计算机程序对比序列信息,进行对比。一种示例性的、优选的计算机程序是遗传学计算机组(Genetic Computer Group, GCG?; Madison, WI) WISCONSIN PACKAGE?10.0版程序, “GAP”(Devereux等人, 1984, Nucl. Acids Res., 12:387)。除了在2个核酸序列之间进行对比以外,该“GAP”程序可以用于在2个氨基酸序列之间的对比,以及核酸序列和氨基酸序列之间的对比。“GAP”程序的优选的默认参数包括:(1)核苷酸的一元对比矩阵(对于同一性,含有1的值;对于不同一性,含有0的值)的GCG?实现,和Gribskov和Burgess, Nucl. Acids Res., 14:6745, 1986的加权的氨基酸对比矩阵,参见Schwartz和Dayhoff, 编, “Atlas of Polypeptide Sequence and Structure,”全国生物医学研究基金会(National Biomedical Research Foundation), 第353-358页, 1979,或其它相当的对比矩阵;(2)对于氨基酸序列,每个间隙的罚分为30,每个间隙中的每个符号的额外罚分为1;或者,对于核苷酸序列,每个间隙的罚分为50,每个间隙中的每个符号的额外罚分为3;(3)对末端间隙没有罚分;和(4)对长间隙没有最大罚分。还可以使用本领域技术人员用于序列对比的其它程序,诸如,例如,BLASTN程序2.2.7版,其可从国立医学图书馆(National Library of Medicine)网站得到进行使用,或WU-BLAST 2.0算法(Advanced Biocomputing, LLC)。另外,BLAST算法使用BLOSUM62氨基酸评分矩阵,可以使用的任选的参数如下:(A)包括过滤器,以掩蔽查询序列的具有低组成复杂性的区段(通过Wootton和Federhen (Computers and Chemistry, 1993)的SEG程序来测定;也参见Wootton和Federhen, 1996, “Analysis of compositionally biased regions in sequence databases,”Methods Enzymol., 266: 554-71),或由短周期性内部重复序列组成的区段(通过Claverie和States (Computers and Chemistry, 1993)的XNU程序来测定),和(B)用于报告相对于数据库序列的匹配的统计显著性阈值,或E-评分(仅仅随机地发现预期的匹配概率,根据Karlin和Altschul, 1990的随机模型;如果归于匹配的统计显著性大于该E-评分阈值,将不报告该匹配);优选的E-评分阈值是0.5,或按照递增的优选次序,为:0.25、0.1、0.05、0.01、0.001、0.0001、1e-5、1e-10、1e-15、1e-20、1e-25、1e-30、1e-40、1e-50、1e-75或1e-100。
本文使用的标准的重组DNA和分子克隆技术是本领域众所周知的,更完整地描述在Sambrook,J.,Fritsch,E.F.和Maniatis,T. Molecular Cloning:A Laboratory Manual;Cold Spring Harbor Laboratory Press:Cold Spring Harbor,1989 (在下文中“Sambrook”)。
现在转至实施方案:
实施方案包括:可用于赋予干旱耐受性的分离的多核苷酸和多肽、重组DNA构建体,包含这些重组DNA构建体的组合物(诸如植物或种子),以及使用这些重组DNA构建体的方法。
分离的多核苷酸和多肽:
本公开内容包括下述的分离的多核苷酸和多肽:
一种分离的多核苷酸,其包含:(i)编码特定多肽的核酸序列,所述多肽具有与SEQ ID NO: 1相比至少50%、51%、52%、53%、54%、55%、56%、57%、58%、59%、60%、61%、62%、63%、64%、65%、66%、67%、68%、69%、70%、71%、72%、73%、74%、75%、76%、77%、78%、79%、80%、81%、82%、83%、84%、85%、86%、87%、88%、89%、90%、91%、92%、93%、94%、95%、96%、97%、98%、99%或100%序列同一性的氨基酸序列,这基于Clustal V比对方法,或(ii) (i)的核酸序列的完整互补体,其中所述完整互补体和(i)的核酸序列由相同数目的核苷酸组成,且是100%互补的。任一种前述分离的多核苷酸可以用于本公开内容的任意重组DNA构建体中。所述多肽优选地是PHD1多肽。
一种分离的多肽,所述多肽具有与SEQ ID NO: 1相比至少50%、51%、52%、53%、54%、55%、56%、57%、58%、59%、60%、61%、62%、63%、64%、65%、66%、67%、68%、69%、70%、71%、72%、73%、74%、75%、76%、77%、78%、79%、80%、81%、82%、83%、84%、85%、86%、87%、88%、89%、90%、91%、92%、93%、94%、95%、96%、97%、98%、99%或100%序列同一性的氨基酸序列,这基于Clustal V比对方法。所述多肽优选地是PHD1多肽。
一种分离的多肽,其中通过至少一种选自缺失、置换、添加和插入的方法,通过改变一个或多个氨基酸,从SEQ ID NO: 1衍生出所述氨基酸序列;和(c)一种多肽,其中所述多肽的氨基酸序列包含SEQ ID NO: 1。所述多肽优选地是PHD1多肽。
一种分离的多核苷酸,其包含编码具有差向异构酶活性的多肽的核苷酸序列,其中所述核苷酸序列可在严谨条件下与包含SEQ ID NO: 18-34的完整互补体的DNA分子杂交。
例如,可以如下完成定位诱变:使用合成的寡核苷酸引物,除了具有特定错配(即,希望的突变)以外,所述引物与要突变的单链噬菌体DNA互补。也就是说,使用上述的合成的寡核苷酸作为引物,以造成噬菌体合成互补链,然后使用得到的双链体DNA转化宿主细胞。将转化的细菌培养物铺板在琼脂上,从而允许从含有噬菌体的单个细胞形成噬斑。作为结果,在理论上,50% 的新菌落含有具有单链突变的噬菌体,而剩下的50%具有原始序列。在特定温度(该温度允许与具有上述希望突变的DNA的完全相同的DNA杂交,但是不允许与具有原始链的DNA杂交),使得到的噬斑与通过激酶处理进行标记的合成探针杂交。随后,挑取与所述探针杂交的噬斑,并培养,用于收集它们的DNA。
术语“在严谨条件下”是指,2个序列在中等或非常严谨的条件下杂交。更具体地,中等严谨的条件可以由本领域普通技术人员容易地确定,例如,取决于DNA的长度。基本条件参见:Sambrook等人, Molecular Cloning: A Laboratory Manual, 第3版, 第6和7章, Cold Spring Harbor Laboratory Press, 2001,且包括使用硝酸纤维素过滤器的预洗溶液5xSSC、0.5% SDS、1.0 mM EDTA (pH 8.0),约50% 甲酰胺、2xSSC至6xSSC在约40-50℃的杂交条件(或其它类似的杂交溶液,诸如Stark氏溶液,在约50% 甲酰胺中,在约42℃)和例如约40-60℃、0.5-6xSSC、0.1% SDS的洗涤条件。优选地,中等严谨的条件包括:在约50℃和6xSSC的杂交(和洗涤)。非常严谨的条件还可以由本领域技术人员容易地确定,例如,取决于DNA的长度。
通常,这样的条件包括,与中等严谨的条件相比,在更高的温度和/或更低的盐浓度杂交和/或洗涤(诸如在约65℃、6xSSC至0.2xSSC、优选地6xSSC、更优选地2xSSC、最优选地0.2xSSC杂交)。例如,非常严谨的条件可以包括:如上定义的杂交,和在大约65-68℃、0.2xSSC、0.1% SDS洗涤。SSPE (1xSSPE是0.15 M NaCl、10 mM NaH2PO4和1.25 mM EDTA,pH 7.4)可以替代在杂交和洗涤缓冲液中的SSC (1xSSC是0.15 M NaCl和15 mM柠檬酸钠);在杂交结束以后,进行洗涤15分钟。
“抑制DNA构建体”是一种重组DNA构建体,当转化或稳定整合入植物的基因组中时,导致植物中靶基因的“沉默”。所述靶基因可以是该植物内源的或转基因的。本文关于靶基因使用的“沉默”,通常是指靶基因表达的mRNA或蛋白/酶的水平的抑制,和/或酶活性水平或蛋白功能性的抑制。术语“抑制”和“沉默”在本文中可互换地使用,包括降低、减少、下降、减退、抑制、消除和阻止。“沉默”或“基因沉默”不特指机制,且是包含性的,不限于,反义、共抑制、病毒抑制、发夹抑制、茎-环抑制、基于RNAi的方案、和基于小RNA的方案。
抑制DNA构建体可以包含源自目标靶基因的区域,且可以包含目标靶基因的有义链(或反义链)的全部或部分核酸序列。根据要使用的方案,所述区域可以与目标基因的全部或部分有义链(或反义链)具有100%同一性或小于100%同一性(例如,至少50%、51%、52%、53%、54%、55%、56%、57%、58%、59%、60%、61%、62%、63%、64%、65%、66%、67%、68%、69%、70%、71%、72%、73%、74%、75%、76%、77%、78%、79%、80%、81%、82%、83%、84%、85%、86%、87%、88%、89%、90%、91%、92%、93%、94%、95%、96%、97%、98%或99%同一性)。
抑制DNA构建体是本领域已知的,一旦选定目标靶基因后可以容易地构建,包括、但不限于,共抑制构建体,反义构建体,病毒抑制构建体,发夹抑制构建体,茎-环抑制构建体,双链RNA-生产构建体,和更一般地,RNAi (RNA干扰)构建体和小RNA构建体例如siRNA (短干扰RNA)构建体和miRNA (微RNA)构建体。
“反义抑制”指能抑制靶基因或基因产物的表达的反义RNA转录物的生产。“反义RNA”是指与全部或部分的靶初级转录物或mRNA互补、且阻断分离的靶核酸片段的表达的RNA转录物(美国专利号5,107,065)。反义RNA的互补性可以是与特定基因转录物的任意部分,即,在5'非编码序列、3'非编码序列、内含子或编码序列处。
“共抑制”是指能抑制靶基因或基因产物的表达的有义RNA转录物的生产。“有义”RNA是指一种RNA转录物,其包括mRNA,且可以在细胞内或体外翻译成蛋白。通过集中在与天然mRNA在有义方向具有同源性的核酸序列的过表达,以前已经设计了植物中的共抑制构建体,它会导致与过表达的序列具有同源性的所有RNA的减少(参见Vaucheret等人, Plant J. 16:651-659 (1998);和Gura, Nature 404:804-808 (2000))。
另一个变化形式描述了植物病毒序列用于指导邻近mRNA编码序列的抑制的应用(PCT公开号WO 98/36083,于1998年8月20日公开)。
调节序列:
本公开内容的重组DNA构建体可以包含至少一个调节序列。调节序列可以是启动子。
许多启动子可以用于本公开内容的重组DNA构建体中。启动子可以基于希望的结果来选择,且可以包括组成型启动子、组织特异性的启动子、诱导型启动子、或用于在宿主生物体中表达的其它启动子。造成基因在大部分时间在大多数细胞类型中表达的启动子通常称作“组成型启动子”。
在35S或UBI启动子控制下的候选基因的高水平组成型表达可以具有多效影响,尽管当由组成型启动子驱动时可以估测候选基因效力。组织特异性的和/或胁迫特异性的启动子的应用,可以消除不希望的影响,但是保留增强干旱耐受性的能力。该影响已经在拟南芥属中观察到(Kasuga等人(1999) Nature Biotechnol. 17:287-91)。
适用于植物宿主细胞中的组成型启动子包括,例如,Rsyn7启动子的核心启动子,和在WO 99/43838和美国专利号6,072,050中公开的其它组成型启动子;核心CaMV 35S启动子(Odell等人,Nature 313:810-812 (1985));稻肌动蛋白(McElroy等人,Plant Cell 2:163-171 (1990));泛素(Christensen等人,Plant Mol. Biol. 12:619-632 (1989)和Christensen等人,Plant Mol. Biol. 18:675-689 (1992));pEMU (Last等人,Theor. Appl. Genet. 81:581-588 (1991));MAS (Velten等人,EMBO J. 3:2723-2730 (1984));ALS启动子(美国专利号5,659,026),等。其它组成型启动子包括,例如,在美国专利号5,608,149、5,608,144、5,604,121、5,569,597、5,466,785、5,399,680、5,268,463、5,608,142和6,177,611中讨论的那些。
在选择用于本公开内容的方法中的启动子时,可能希望使用组织特异性的或发育调节的启动子。组织特异性的或发育调节的启动子是DNA序列,其调节DNA序列在对于雄花穗发育、结实、或二者的发育至关重要的植物细胞/组织中的选择性表达,并将这样的DNA序列的表达限制在植物的雄花穗发育或种子成熟的阶段。任意可识别的启动子可以用于本公开内容的方法中,造成希望的时间和空间表达。叶子特异性的启动子是合适的。
响应于内源或外源刺激例如化学化合物(化学诱导物)的存在,或响应于环境的、激素的、化学的、和/或发育的信号,诱导型启动子选择性地表达可操作地连接的DNA序列。诱导型或调节的启动子包括,例如,由光、热、胁迫、水灾或干旱、植物激素、创伤或化学物质(例如乙醇、茉莉酮酸酯、水杨酸或安全剂)调节的启动子。
使用的启动子包括下述的:1)胁迫可诱导的RD29A启动子(Kasuga等人(1999) Nature Biotechnol. 17:287-91);2)大麦启动子,B22E;在发育中的玉米核中,B22E的表达是梗特异性的(“Primary Structure of a Novel Barley Gene Differentially Expressed in Immature Aleurone Layers”. Klemsdal, S.S. 等人, Mol. Gen. Genet. 228(1/2):9-16 (1991));和3)玉米启动子,Zag2 (“Identification and molecular characterization of ZAG1, the maize homolog of the Arabidopsis floral homeotic gene AGAMOUS”, Schmidt, R.J. 等人, Plant Cell 5(7):729-737 (1993);“Structural characterization, chromosomal localization and phylogenetic evaluation of two pairs of AGAMOUS-like MADS-box genes from maize”, Theissen等人Gene 156(2):155-166 (1995); NCBI GenBank登记号X80206))。Zag2转录物在授粉前5天至授粉后7-8天(授粉后天数,“DAP”)可以检测到,并指导在发育中的雌花序的心皮中的表达,和对发育中的玉米仁的核特异性的Ciml。Ciml转录物在授粉前4-5天至授粉后6-8天可以检测到。其它有用的启动子包括可以源自其表达在母系上与发育中的雌小花有关的基因的任意启动子。
启动子可以整体源自天然基因,或可以由在自然界发现的不同启动子所衍生出的不同元件组成,或甚至包含合成的DNA区段。
本公开内容的重组DNA构建体还可以包括其它调节序列,包括、但不限于, 翻译前导序列、内含子和聚腺苷酸化识别序列。在本公开内容的另一个实施方案中,本公开内容的重组DNA构建体另外包含增强子或沉默子。
可以将内含子序列添加到5’非翻译区、蛋白编码区或3’非翻译区上,以增加在胞质溶胶中积累的成熟信号的量。已经证实,在植物和动物表达构建体的转录单元中包含可剪接的内含子,会在mRNA和蛋白水平增加基因表达最高达1000倍。Buchman和Berg, Mol. Cell Biol. 8:4395-4405 (1988); Callis等人, Genes Dev. 1:1183-1200 (1987)。
可以选择任意的植物,用于鉴别将用于本公开内容的重组DNA构建体中的调节序列和PHD1多肽基因。适用于分离基因和调节序列的植物靶标的实例包括、但不限于:苜蓿、苹果、杏、拟南芥、朝鲜蓟、芝麻菜(arugula)、芦笋、酪梨、香蕉、大麦、菜豆、甜菜、黑莓、蓝莓、绿花椰菜、抱子甘蓝、卷心菜、芸苔、香瓜、胡萝卜、木薯、蓖麻子、花椰菜、芹菜、樱桃、菊苣、芫荽、柑桔、金钱橘、三叶草、椰子、咖啡、玉米、棉花、酸果蔓、黄瓜、花旗松(Douglas fir)、茄子、菊苣、茅菜(escarole)、桉树、茴香、无花果、大蒜、葫芦、葡萄、葡萄柚、蜜汁、豆薯、猕猴桃、莴苣、韭、柠檬、莱姆(lime)、火炬松、亚麻子、芒果、甜瓜、蘑菇、油桃、坚果、燕麦、油椰、油籽油菜、秋葵、橄榄、洋葱、桔子、观赏植物、棕榈、木瓜、荷兰芹、荷兰防风草、豌豆、桃子、花生、梨、胡椒、柿、松、凤梨、车前草、洋李、石榴、白杨、马铃薯、西葫芦、柑橘、放射松、radiscchio、小红萝卜、油菜籽、覆盆子、稻、黑麦、高梁、南方松、大豆、菠菜、南瓜、草莓、甜菜、甘蔗、向日葵、甘薯、香枫、红橘、茶树、烟草、番茄、小黑麦(triticale)、草皮、芜菁、藤、西瓜、小麦、薯芋和胡瓜。
组合物:
本公开内容的组合物是植物,所述植物在它的基因组中包含本公开内容的任意重组DNA构建体(例如上面讨论的任意构建体)。组合物也包括植物的任意后代、和从植物或其后代得到的任意种子,其中所述后代或种子在它的基因组内包含重组DNA构建体。后代包括通过植物的自花授粉或异系杂交得到的后代。后代也包括杂交和近交系。
在杂交种子繁殖的作物中,成熟的转基因植物可以自交,以生产纯合的近交植物。近交植物会生成含有新导入的重组DNA构建体的种子。可以培育这些种子,以生成表现出改变的农艺学特性(例如,增加的农艺学特性,任选地,在水限制条件下)的植物,或用于育种程序,以生产杂交种子,后者可以生长,以生成表现出这样的改变的农艺学特性的植物。所述种子可以是玉米种子。
所述植物可以是单子叶的或双子叶植物,例如,玉米、稻或大豆植物,诸如玉米杂交植物或玉米近交植物。所述植物也可以是向日葵、高粱、油菜、小麦、苜蓿、棉花、大麦、粟、甘蔗、柳枝稷、烟草、马铃薯和糖用甜菜。
所述重组DNA构建体可以稳定地整合进植物的基因组中。
所述PHD1多肽可以来自拟南芥(Arabidopsis thaliana)、玉米(Zea mays)、大豆(Glycine max)、烟豆(Glycine tabacina)、劳豆(Glycine soja)或短绒野大豆(Glycine tomentella)。
至少一种农艺学特性可以选自:绿度、产量、生长速率、生物量、成熟鲜重、成熟干重、果实产量、种子产量、总植物氮含量、果实氮含量、种子氮含量、营养组织中的氮含量、总植物游离氨基酸含量、果实游离氨基酸含量、种子游离氨基酸含量、营养组织中的游离氨基酸含量、总植物蛋白含量、果实蛋白含量、种子蛋白含量、营养组织中的蛋白含量、干旱耐受性、氮吸收、根倒伏、收获指数、茎倒伏、植物高度、穗高度、穗长度、盐耐受性、早期幼苗活力和在低温胁迫下的出苗。例如,所述至少一种农艺学特性的改变可以是产量增加、绿度或生物量。
在本公开内容的前述实施方案1-8中的任一个或任意其它实施方案中,在水限制条件下,与不包含所述重组DNA构建体的对照植物相比,所述植物可以表现出至少一种农艺学特性的改变。
“干旱”是指植物的水可用性的减少,特别当持续长时间时,可以造成对植物的损伤或阻止它的成功生长(例如,限制植物生长或种子产量)。
“干旱耐受性”是植物在干旱条件下存活长时间段且不表现出实质的生理学或植株恶化的特性。
植物的“增加的干旱耐受性”相对于参照或对照植物来测量,且是这样的植物特性:所述植物在干旱条件下存活长时间段,且不表现出与在类似的干旱条件下生长的参照或对照植物相同程度的生理学或植株恶化。通常,当在它的基因组中包含重组DNA构建体的转基因植物表现出与参照或对照植物相比增加的干旱耐受性时,所述参照或对照植物在它的基因组中不包含所述重组DNA构建体。
本领域普通技术人员熟知:用于模拟干旱条件的方案,和用于评价已经遭受模拟的或天然存在的干旱条件的植物的干旱耐受性的方案。例如,通过在一段时间内给植物提供比正常所需更少的水或不提供水,可以模拟干旱条件,通过寻找生理学和/或植株状况的差异,可以评价干旱耐受性,所述状况包括(但不限于)活力、生长、大小或根长度,或具体地,叶颜色或叶面积大小。用于评价干旱耐受性的其它技术包括:测量叶绿素荧光、光合速率和气体交换速率。
干旱胁迫实验可以包括慢性胁迫(即,缓慢地枯干),和/或可以包括被1天或2次恢复隔开的2次急性胁迫(即,突然除去水)。慢性胁迫可以持续8-10天。急性胁迫可以持续3-5天。在转基因植物和相关对照植物的干旱胁迫和良好给水处理的过程中,可以测量下述变量:
变量“% area chg_start chronic - acute2”是通过远可见光谱成像测得的慢性胁迫第一天和第二次急性胁迫当天之间的总面积的变化百分比的量度。
变量“% area chg_start chronic - end chronic”是通过远可见光谱成像测得的慢性胁迫第一天和慢性胁迫最后一天之间的总面积的变化百分比的量度。
变量“% area chg_start chronic - harvest”是通过远可见光谱成像测得的慢性胁迫第一天和收获当天之间的总面积的变化百分比的量度。
变量“% area chg_start chronic - recovery24hr”是通过远可见光谱成像测得的慢性胁迫第一天和恢复24小时(在急性胁迫2以后24小时)之间的总面积的变化百分比的量度。
变量“psii_acute1”是在第一次急性胁迫期结束时光系统II (PSII)效率的量度。它会提供PSII天线吸收光的效率的估测,且与叶内的二氧化碳同化直接相关。
变量“psii_acute2”是在第二次急性胁迫期结束时光系统II (PSII)效率的量度。它会提供PSII天线吸收光的效率的估测,且与叶内的二氧化碳同化直接相关。
变量“fv/fm_acute1”是在第一次急性胁迫结束时最适量子产率(Fv/Fm)(最大和最小荧光之间的可变荧光差异/最大荧光)的量度。
变量“fv/fm_acute2”是在第二次急性胁迫结束时最适量子产率(Fv/Fm)(最大和最小荧光之间的可变荧光差异/最大荧光)的量度。
变量“leaf rolling_harvest”是在收获当天上图与侧图之比的量度。
变量“leaf rolling_recovery24hr”是在恢复24小时时上图与侧图之比的量度。
变量“比生长速率(SGR)”描述了总植物表面积(通过Lemna Tec Instrument测得)在一天中的变化(Y(t) = Y0*er*t )。Y(t) = Y0*er*t等同于Y/Δt的变化百分比,其中各个术语如下:Y(t) = 在t时的总表面积;Y0 = 最初的总表面积(估测);r = 比生长速率/天,且t =种植以后的天数(“DAP”)。
变量“茎干干重”是放入104℃烘箱中以后96小时的茎干重量的量度。
变量“茎干鲜重”是刚从植物切下以后的茎干重量的量度。
下面的实施例描述了用于模拟干旱条件和/或评价干旱耐受性的一些代表性的方案和技术。
方法包括、但不限于下述的:
一种用于转化细胞的方法,所述方法包括:用本公开内容的任一种分离的多核苷酸转化细胞。也包括通过该方法转化的细胞。在具体实施方案中,所述细胞是真核细胞,例如,酵母、昆虫或植物细胞,或原核细胞,例如,细菌细胞。
一种用于生产转基因植物的方法,所述方法包括:用本公开内容的任一种分离的多核苷酸或重组DNA构建体转化植物细胞,和从所述转化的植物细胞再生转基因植物。本公开内容也涉及通过该方法生产的转基因植物和从该转基因植物得到的转基因种子。
一种用于从细胞或细胞培养基分离本公开内容的多肽的方法,其中所述细胞包含重组DNA构建体,所述构建体包含与至少一个调节序列可操作地连接的本公开内容的多核苷酸,且其中在适合表达所述重组DNA构建体的条件下,培养转化的宿主细胞。
一种改变本公开内容的多肽在宿主细胞中的表达水平的方法,所述方法包括:(a)用本公开内容的重组DNA构建体转化宿主细胞;和(b)在适合表达所述重组DNA构建体的条件下,培养所述转化的宿主细胞,其中所述重组DNA构建体的表达会导致在所述转化的宿主细胞中生产改变水平的本公开内容的多肽。
一种增加植物的产量和/或干旱耐受性的方法,所述方法包括:(a)向可再生的植物细胞中导入重组DNA构建体,所述构建体包含与至少一个调节序列(例如,在植物中有功能的启动子)可操作地连接的多核苷酸,其中所述多核苷酸编码的多肽具有这样的氨基酸序列,所述氨基酸序列与SEQ ID NO: 1相比具有至少50%、51%、52%、53%、54%、55%、56%、57%、58%、59%、60%、61%、62%、63%、64%、65%、66%、67%、68%、69%、70%、71%、72%、73%、74%、75%、76%、77%、78%、79%、80%、81%、82%、83%、84%、85%、86%、87%、88%、89%、90%、91%、92%、93%、94%、95%、96%、97%、98%、99%或100%序列同一性,这基于Clustal V比对方法,和(b)从步骤(a)以后的可再生的植物细胞再生转基因植物,其中所述转基因植物在它的基因组中包含所述重组DNA构建体,并表现出与不包含所述重组DNA构建体的对照植物相比增加的干旱耐受性。所述方法可以另外包括:(c)获得从所述转基因植物衍生出的后代植物,其中所述后代植物在它的基因组中包含所述重组DNA构建体,并表现出与不包含所述重组DNA构建体的对照植物相比增加的干旱耐受性。
实施例
在下面的实施例中进一步解释了本公开内容,其中份数和百分比按重量计,度数是摄氏度,除非另有说明。应当理解,这些实施例尽管指明了本公开内容的优选实施方案,仅仅作为解释而给出。从上面的讨论和这些实施例中,本领域技术人员可以确定本公开内容的基本特征,在不背离其精神和范围的情况下,可以作出本公开内容的多种变化和改进,使它适用于多种用途和条件。因而,本领域技术人员从前面的描述,可以明白除了本文显示和描述的那些以外的本公开内容的多种改进。这样的改进也在所附权利要求书的范围内。
实施例1
phd1突变体的分离和表征
在具有改变的营养器官中的碳水化合物水平的480个突变体系中,选择具有低碳水化合物含量和生长矮小的光同化物缺陷型1 (phd1)用于进一步表征(图1)。与野生型相比,在光照/黑暗周期中设定的所有时间点,所述突变体的成熟叶子具有低淀粉水平(图1I)。秆的扫描电子显微照片证实,在phd1突变体的薄壁组织细胞中沉积更少的淀粉颗粒。在幼苗期中,突变体的茎干和初生根比野生型的那些更短和更轻(图1A)。在节间伸长以后,phd1突变体表现出半矮生、更少的分蘖、迟缓的营养生长、更晚的开花和更少的籽粒灌浆表型(图1B-E)。另外,尽管每穗粒数在突变体和野生型之间没有变化,phd1突变体的结籽率显著降低,这最终导致籽粒产量的显著下降(图1F、G)。
实施例2
PHD1编码功能性的叶绿体UDP-Glc差向异构酶
遗传分析指示,phd1表型由单个隐性基因控制,所述隐性基因不会与T-DNA插入物共分离,因此进行了基于图谱的克隆。在物理图谱中将PHD1基因座划界至染色体1的短臂上的72-kb区域。该区域含有6个注解基因,对来自phd1-1的这些基因测序,鉴别出在Os01g0367100的外显子2中的单个核苷酸转换(G至T),这会导致过早的翻译终止。通过分析2个其它的phd1等位基因(分离自相同的遗传筛选),证实了Os01g0367100作为PHD1的身份。发现了在phd1-2的外显子7中的单个核苷酸置换(A至T)和在phd1-3的外显子3和外显子4之间的13-bp插入物(图2A)。在3个等位基因突变体中的任一个中,都几乎没有检测到PHD1 mRNA (图S2)。通过野生型Os01g0367100在phd1-1突变体背景中的转基因表达,完全补足了phd1表型(图2B、C),这证实,Os01g0367100的无义突变会引起假定的无效突变体表型。
PHD1与得自假微型海链藻(Thalassiosira pseudonana)(XP_002290295)、三角褐指藻(Phaeodactylum tricornutum)(XP_002178225)、莱茵衣藻(Chlamydomonas reinhardtii)(XP_001699105)、Micromonas pusilla (EEH60780)、Ostreococcus tauri (CAL54696)、小立碗藓(Physcomitrella patens)(XP_001767242)、蓖麻(XP_002516868)、拟南芥(AT2G39080)、毛果杨(XP_002311843)、葡萄(XP_002276706)、玉米(NP_001131736)和高粱(XP_002457832)的蛋白(通过引用并入本文)具有相似性,具有27-75%的氨基酸同一性(图S3)。在PHD1和它的16种结构上类似的蛋白之间的系统进化分析表明,PHD1与得自高粱的Sb03g014730和得自玉米的LOC100193101密切相关(图3)。仅在植物界中发现了PHD1同系物/直系同源物,这表明,这些蛋白在植物物种之间是进化上保守的。保守结构域的分析证实,PHD1和它的同系物含有共有的WcaG结构域,这是核苷-二磷酸糖差向异构酶的特征(图11A)。UDP-Glc差向异构酶(UGE)(一种糖差向异构酶)会催化UDP-Glc和UDP-Gal的互变。PHD1和它的同系物可以起新颖的植物特异性的UGE的作用。
为了理解PHD1的作为UDP-Glc差向异构酶的生化功能,在大肠杆菌中表达了缺少假定的N-端62个氨基酸转运肽的成熟PHD1蛋白,并检查了UGE活性。结果表明,PHD1可以催化UDP-Gal向UDP-Glc的转化,且曲线拟合指示,UDP-Gal结合符合简单的米-门二氏动力学,在30℃时具有0.84 mM的K m值(图12A)。为了检查PHD1在体内是否具有UDP-Glc差向异构酶活性,从酵母甘油醛-3-磷酸脱氢酶启动子表达成熟的PHD1,以补足酵母gal10Δ突变体(其不能在含有D-半乳糖作为唯一碳源的培养基上生长)的营养缺陷表型。补足结果证实,PHD1在体内也具有UDP-Glc差向异构酶活性(图12B)。
RNA凝胶印迹分析揭示,PHD1存在于所有绿色组织中,在叶片和叶鞘中最丰富,其次是花和秆,但是仅以非常低的水平存在于根中(图4A)。mRNA原位杂交揭示,PHD1主要在叶原基和茎干顶端分生组织(图4B)、嫩叶的围绕维管束的叶肉细胞(图4C)、花序原基(图4D)和腋芽(图4E)中表达。PHD1编码340个氨基酸的蛋白,该蛋白具有在N-端处的假定的62个氨基酸的叶绿体转运肽。为了证实PHD1的叶绿体定位,使全长PHD1与在花椰菜花叶病毒(CaMV) 35S启动子控制下的绿荧光蛋白(GFP)报道基因融合,随后转化进稻茎干原生质体中。图4F表明,GFP荧光与红叶绿素自发荧光共局部化,这证实,PHD1是定位于叶绿体的蛋白,且预测的转运肽是有功能的。
实施例3
PHD1会影响叶子中的核苷酸糖和碳同化的体内稳态
UDP-Gal是生物合成反应中的活化形式的半乳糖,但是半乳糖补救途径存在于真核生物中。为了评估在Leloir补救途径中涉及的基因的表达,分析了该途径的3个关键基因GalM、GalK和GalT在phd1-1和野生型中的表达水平。所有3个基因的表达在phd1-1突变体中都被显著地增量调节,这提示整个补救途径的活化(图5A)。β-乳糖酶参与从多糖分解来产生游离的β-D-Gal,且UDP-Glc焦磷酸化酶(UGP)会催化从Glc-1-P形成UDP-Glc。编码β-乳糖酶和UGP3的基因的表达水平也在phd1-1中被增量调节。令人惊奇地,在稻中编码UGE的推定细胞质同种型的OsUGE1和OsUGE4在phd1-1中的表达水平比在野生型植物中高了超过2倍,这指示,细胞质中从头UDP-Gal生物合成的增量调节。这些结果提示,PHD1可能负责叶绿体中的大部分UGE功能,且似乎参与从UDP-Glc产生UDP-Gal,以供给用于半乳糖脂生物合成(为适当的叶绿体膜组成所需)的结构单元。
为了确定在phd1-1中的UGE活性的损害是否对含有半乳糖的化合物具有影响,分析了半乳糖代谢的中间体。发现UDP-Gal在phd1-1的叶子中的量稍微高于在野生型中,但是UDP-Glc量显著更高(图6A)。因此,在phd1-1中的UDP-Glc与UDP-Gal之比也高于在野生型叶子中。这些结果提示,PHD1功能障碍可以触发底物的积累,并扰乱2种糖核苷酸之间的互变平衡。
因为phd1-1突变体表现出碳同化物水平的显著降低,测定了在淀粉和蔗糖的合成、运输和裂解中涉及的几个关键基因的转录物水平在野生型和phd1-1植物的成熟叶子中是否改变。令人感兴趣地,尽管参与淀粉生物合成的基因(诸如AGPL2、SSI、SSIIIa、GBSS、BE和BT1)的表达在phd1-1突变体中受到抑制(图5B),参与蔗糖裂解的基因(诸如INV1/3和SuSy1)的表达水平都增加(图5C)。同时,编码葡萄糖-6-磷酸/磷酸盐转运蛋白的GPT基因在phd1-1中被增量调节,这指示己糖-磷酸从叶绿体输出至胞质溶胶的增强。另外,增加的UGP2(一个参与UDP-Glc合成的基因)表达水平与phd1-1突变体中增加的UDP-Glc积累和更高的UDP-Glc/UDP-Gal比率相关。
实施例4
PHD1功能障碍会影响光合膜系统
叶绿体膜含有高水平的糖脂,UDP-Gal是糖脂生物合成的优势底物。为了检查PHD1功能障碍对膜脂质体内稳态的影响,分析了从phd1-1、野生型和PHD1-补足的植物提取的总脂质的组成(图6B)。在phd1-1突变体中,MGDG的mol% 量与野生型和补足的植物相比降低了19%,这指示,PHD1参与MGDG生物合成。相比而言,在phd1-1突变体中仅观察到DGDG含量的轻微下降(2.5%),这证实,DGDG合成可以不需要PHD1,并提示,用于形成DGDG的UDP-Gal底物可能从胞质溶胶供给。在phd1-1中的MGDG的丰度的降低,伴随着其它重要膜脂质(诸如磷脂酰胆碱(PC)、磷酸肌醇(PI)和磷脂酰甘油(PG))的丰度的增加,而磺基奎诺糖基二酰基甘油(SQDG)和磷脂酰乙醇胺(PE)在phd1-1突变体中的mol%水平仅仅轻微地、但是不显著地更高(图6B)。因为PC、PI和PG也是类囊体膜的组分,这些结果证实,突变体类囊体膜具有增加的磷脂的mol% 量。
非侵入性的叶绿素荧光测量指示,phd1-1和野生型具有类似的光系统II光化学的最大量子产率(F v/F m) (表1)。在突变体中,在光系统II中的光化学能量转换的有效量子产率(ΦPSII)轻微地、但是显著地下降(表1)。在phd1-1突变体中的色素含量也下降(表1)。令人感兴趣地,在2月龄植物中,在phd1-1突变体中的叶绿体显著小于野生型植物(野生型,5.0±0.4 μm;phd1-1,3.0±0.5 μm),在突变体中的淀粉颗粒也是要么不存在,要么具有减小的尺寸和/或数目。这些数据指示,在叶绿体中减少的半乳糖脂的量可能降低高等植物的光合能力。
表1. 野生型、phd1-1和PHD1-补足的植物的色素含量(mg·g-1鲜重)和光合参数
从在水稻田中生长的4月龄植物的完全展开的叶子收集样品。值代表6-10个独立测定的平均值(±标准差)。* 突变体和野生型之间的显著差异(P < 0.05)。
实施例5
PHD1的表达会增加生长速率和籽粒产量
研究了PHD1过表达是否会提高生物量和籽粒产量。当在水稻田中生长时,过表达PHD1的转基因稻植物表现出分蘖(分枝)速率和光合速率的显著增加(图7A、表2)。转基因植物的生长速率在幼苗期加速,在成熟期时的干物质积累比非转基因植物增加了12.5%至22.4% (图7D、表2)。另外,与非转基因对照植物相比,转基因系S3、S5和S8的每株籽粒产量分别增加了10.7、15.5和18.3%(图7C)。这些结果证实,在稻中的PHD1过表达与生物量生产和籽粒产量的增加正相关。
表2. 过表达PHD1的植物和野生型的生物量和光合速率的表征(Nipponbare).
S3、S5和S8代表3个独立的过表达PHD1的转基因系。a20天龄稻幼苗;bNPR,在抽穗期测量的净光合速率( μmol·CO2·m-2·s-1);值是得自至少30个植物/系的平均值±标准差,* 显著差异(P < 0.05)。
表3. 用于半乳糖代谢分析的寡核苷酸(SEQ ID NO: 35-52, 连续的)
表4. 用于碳水化合物代谢分析的寡核苷酸(SEQ ID NO: 53-76, 连续的)
实施例9
根瘤土壤杆菌
LBA4404的电穿孔
在冰上融化(20-30 min)电穿孔感受态细胞(40 μL),诸如含有PHP10523的根瘤土壤杆菌(Agrobacterium tumefaciens)LBA4404 (“pSB1”; Komari等人, Plant J. 10:165-174 (1996); NCBI General Identifier No. 59797027)。PHP10523含有:用于T-DNA转移的VIR基因、土壤杆菌低拷贝数质粒复制起点、四环素抗性基因和用于体内DNA双分子重组的Cos位点。同时,在冰上冷冻电穿孔容器。将电穿孔或设置调节至2.1 kV。仍然在冰上,将DNA等分试样(0.5μL亲本DNA,浓度为0.2μg -1.0μg,在低盐缓冲液或双蒸馏水中)与融化的根瘤土壤杆菌LBA4404细胞混合。将所述混合物转移至电穿孔容器的底部,并在冰上静置1-2 min。通过按下“脉冲”按钮2次(理论上,实现4.0毫秒脉冲),将细胞电穿孔(Eppendorf电穿孔仪2510)。随后,将0.5 mL室温2xYT培养基(或SOC培养基)加入该容器中,并转移至15 mL按盖试管(例如,Falcon?试管)。将所述细胞在28-30℃、200-250 rpm温育3 h。
将250 μL等分试样铺展在含有YM培养基和50μg/mL壮观霉素的平板上,并在28-30℃温育3天。为了增加转化体的数目,进行下述2个任选的步骤之一:
选项1: 用30 μL 15 mg/mL利福平覆盖平板。LBA4404具有利福平的染色体抗性基因。该额外选择会消除当使用LBA4404感受态细胞的更差制品时观察到的一些污染菌落。
选项2: 执行2次电穿孔重复,以补偿更差的电感受态细胞。
转化体的鉴别:
挑选4个独立菌落,并在含有AB基本培养基和50μg/mL壮观霉素的平板上划线,用于分离单个菌落。将所述平板在28℃温育2-3天。挑选每个假定的共合体的单个菌落,并接种4 mL 10 g/L细菌用蛋白胨、10 g/L酵母浸出物、5 g/L氯化钠和50 mg/L壮观霉素。在摇动下,将所述混合物在28℃温育24 h。使用QIAGEN?Miniprep和任选的Buffer PB洗液,从4 mL培养物中分离出质粒DNA。在30 μL中洗脱所述DNA。如上所述,将2 μL等分试样用于电穿孔20 μL DH10b + 20 μL双蒸馏水。任选地,可以使用15 μL等分试样转化75-100 μL Invitrogen?Library Efficiency DH5α。将所述细胞铺展在含有LB培养基和50μg/mL壮观霉素的平板上,并在37℃温育过夜。
挑选每个假定的共合体的3-4个独立菌落,并接种4 mL含有50 μg/mL壮观霉素的2xYT培养基(10 g/L细菌用蛋白胨、10 g/L酵母浸出物、5 g/L氯化钠)。在摇动下,将所述细胞在37℃温育过夜。接着,使用QIAPREP?Miniprep和任选的Buffer PB洗液,从4 mL培养物中分离出质粒DNA(在50 μL中洗脱)。使用8 μL用于SalI消化(使用亲本DNA和PHP10523作为对照)。对于4个代表2种具有正确SalI消化模式的假定共合体的质粒,进行另外3种使用限制性酶BamHI、EcoRI和HindIII的消化(使用亲本DNA和PHP10523作为对照)。推荐电凝胶用于对比。
实施例11
使用土壤杆菌转化玉米
基本上如Zhao等人在Meth. Mol. Biol. 318:315-323 (2006) (也参见Zhao等人, Mol. Breed. 8:323-333 (2001)和1999年11月9日授权的美国专利号5,981,840,通过引用并入本文)所述,进行土壤杆菌介导的玉米转化。所述转化过程包括细菌接种、共培养、静止(resting)、选择和植物再生。
1. 未成熟胚制备:
从颖果切下未成熟的玉米胚,并放入含有2 mL PHI-A培养基的2 mL微管中。
2. 未成熟胚的土壤杆菌感染和共培养:
2.1感染步骤:
用1 mL微量移液器取出(1)的PHI-A培养基,并加入1 mL土壤杆菌悬浮液。轻轻倒转所述试管进行混合。将所述混合物在室温温育5 min。
2.2共培养步骤:
使用1 mL微量移液器,取出得自感染步骤的土壤杆菌悬浮液。使用无菌铲,从试管刮下胚,并转移至在100x15 mm培养皿中的PHI-B培养基平板。使胚轴向下,将所述胚定向在所述培养基的表面上。在20℃在黑暗中培养含有胚的平板3天。在共培养期可以使用L-半胱氨酸。使用标准的二元载体,补充了100-400 mg/L L-半胱氨酸的共培养培养基对于恢复稳定的转基因事件而言是关键性的。
3. 假定的转基因事件的选择:
向在100x15 mm培养皿中的每个PHI-D培养基平板中,转移10个胚,维持朝向,并用PARAFILM?密封所述培养皿。在黑暗中在28℃温育所述平板。预期在6-8周内观察到活跃生长的假定事件,作为淡黄色胚组织。没有发生事件的胚可以是褐色的和坏死的,且几乎不会看到脆性组织生长。根据生长速率,以2-3周间隔,将假定的转基因胚组织传代培养至新鲜的PHI-D平板。记录事件。
4. T0植物的再生:
将在PHI-D培养基上繁殖的胚组织传代培养至在100x25 mm培养皿中的PHI-E培养基(体细胞胚成熟培养基),并在黑暗中在28℃温育约10-18天,直到体细胞胚成熟。将单个的、成熟的体细胞胚(具有边界清楚的胚鳞和胚芽鞘)转移至PHI-F胚萌发培养基,并28℃在光照(约80μE,来自冷白光灯或等效的荧光灯)下温育。在7-10天中,将再生的植物(约10 cm高)盆栽在园艺混合物中,并使用标准的园艺方法稳定化。
用于植物转化的培养基:
1. PHI-A: 4g/L CHU基础盐, 1.0 mL/L 1000X Eriksson氏维生素混合物, 0.5 mg/L盐酸硫胺, 1.5 mg/L 2,4-D, 0.69 g/L L-脯氨酸, 68.5 g/L蔗糖, 36 g/L葡萄糖, pH 5.2。加入100μM乙酰丁香酮(过滤除菌)。
2. PHI-B: PHI-A,不含葡萄糖,将2,4-D增加至2 mg/L,将蔗糖减少至30 g/L,并补充0.85 mg/L硝酸银(过滤除菌)、3.0 g/L GELRITE?、100μM乙酰丁香酮(过滤除菌),pH 5.8。
3. PHI-C: PHI-B,不含有GELRITE?和乙酰丁香酮,将2,4-D减少至1.5 mg/L,并补充8.0 g/L琼脂、0.5 g/L 2-[N-吗啉代]乙烷-磺酸(MES)缓冲液、100 mg/L羧苄西林(过滤除菌)。
4. PHI-D: 补充了3 mg/L双丙氨磷的PHI-C (过滤除菌)。
5. PHI-E: 4.3 g/L的Murashige和Skoog (MS)盐(Gibco, BRL 11117-074), 0.5 mg/L烟酸, 0.1 mg/L盐酸硫胺素, 0.5 mg/L盐酸吡多辛, 2.0 mg/L甘氨酸, 0.1 g/L肌醇, 0.5 mg/L玉米素(Sigma, 目录号Z-0164), 1 mg/L醋酸吲哚(IAA), 26.4μg/L脱落酸(ABA), 60 g/L蔗糖, 3 mg/L双丙氨磷(过滤除菌), 100 mg/L羧苄西林(过滤除菌), 8 g/L琼脂, pH 5.6。
6. PHI-F: PHI-E,不含玉米素、IAA、ABA;将蔗糖减少至40 g/L;用1.5 g/L GELRITE?替代琼脂;pH 5.6。
通过首先将组织簇转移至补充了0.2 mg/升的2,4-D的N6培养基,可以从转基因愈伤组织再生植物。在2周以后,可以将所述组织转移至再生培养基(Fromm等人, Bio/Technology 8:833-839 (1990))。
可以再生转基因T0植物,并测定它们的表型。可以收集T1种子。可以栽培T1植物和/或它们的后代,并测定它们的表型。
实施例15
用PHD1基因转化的玉米系的产量分析
通过直接转化或从分别转化的系的渐渗杂交,可以将含有PHD1基因的重组DNA构建体导入原种玉米近交系中。
转基因植物(近交的或杂交的)可以进行更强烈的基于田地的实验,以在较好供水和水限制条件下研究产量增加和/或稳定性。
可以进行随后的产量分析,以确定与不含有经过验证的耐旱引导基因的对照植物相比,含有经过验证的耐旱引导基因的植物在水限制条件下是否具有产量性状的提高。具体地,可以在含有经过验证的耐旱引导基因的植物和对照植物的开花期和/或灌浆期,施加干旱条件。可以测量二者的产量下降。含有经过验证的耐旱引导基因的植物在水限制条件下具有与对照植物相比更少的产量损失,例如,减少了至少25%的产量损失,或在水不限制条件下,具有与对照植物相比增加的产量。
上述方法可以用于选择这样的转基因植物:其在水限制条件和/或较好供水的条件下,与不包含所述重组DNA构建体的对照植物相比,具有增加的产量。
序列表
<110> Institute of Genetics and Developmental Biology,Chinese Academy of Sciences
Chu, Chengcai
Li, Chunlai
<120> 质体核苷酸糖差向异构酶
<130> RTS16139 -PCT
<150> CN 201010178405.6
<151> 2010-05-17
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260 265 270
Asp Asn Lys Pro Leu Ser Arg Gln Glu Ile Met Asp Ala Val Asn Asn
275 280 285
Ser Gly Lys Phe Asp Thr Lys Phe Gly Gly Phe Thr Gly Thr Asp Gly
290 295 300
Pro Leu Gly Lys Arg Met Glu Asn Ser Lys Thr Arg Ala Glu Ile Gly
305 310 315 320
Trp Glu Pro Lys Tyr Pro Ser Phe Thr Glu Phe Leu Gly Ile Ser Ser
325 330 335
<210> 5
<211> 359
<212> PRT
<213> 玉米
<400> 5
Met Gly Arg Ala Ala Ala Pro Leu Leu Pro Arg Pro Ile Ile Ala Gln
1 5 10 15
Phe His Leu His Pro His His Leu Val His Gln Ser Cys Leu Pro Tyr
20 25 30
Leu His Ala Thr Ala Pro Val Ala Ala Ala Ser Ala Leu Phe Ala Pro
35 40 45
Gly Pro Thr Ser Ser Pro Ser Val Arg Val Ser Arg Pro Arg Arg His
50 55 60
Ala Ser Met Ser Ala Ala Ala Asp Ser Ser Ser Ser Ile Val Val Ser
65 70 75 80
Gly Asp Ala Ala Val Ser Ser Leu Ser Pro Glu Ser Ile Glu His Asn
85 90 95
Asp Leu Leu Ile Val Gly Pro Gly Val Leu Gly Arg Ile Val Ala Glu
100 105 110
Met Trp Lys Gln Glu Tyr Pro Gly Cys Lys Val Tyr Gly Gln Thr Ala
115 120 125
Thr Thr Asp His His Ser Glu Leu Thr Asp Ile Gly Ile Ile Pro Ser
130 135 140
Leu Lys Gly Ser Val Pro Gly Pro Lys Phe Pro Tyr Val Ile Phe Cys
145 150 155 160
Ala Pro Pro Tyr Arg Ser Glu Asp Tyr Ala Gly Asp Leu Arg Val Ala
165 170 175
Ala Ser Asn Trp Asn Gly Lys Gly Ser Phe Leu Phe Thr Ser Ser Thr
180 185 190
Ala Val Tyr Asp Cys Ser Asp Asn Gly Phe Cys Ser Glu Asp Ser Pro
195 200 205
Cys Val Pro Ile Gly Gln Ser Thr Arg Thr Asp Val Leu Leu Lys Ala
210 215 220
Glu Asn Val Val Leu Glu Ala Gly Gly Cys Val Leu Arg Leu Val Gly
225 230 235 240
Leu Tyr Lys Ser Asp Arg Gly Pro His Val Tyr Trp Leu Ser Lys Gly
245 250 255
Thr Leu Asp Val Arg Pro Asp His Ile Leu Asn Leu Ile His Tyr Glu
260 265 270
Asp Ala Ala Ser Leu Val Ile Ser Ile Met Lys Lys Arg Leu Arg Ser
275 280 285
Cys Ile Phe Val Gly Cys Asp Asn Glu Pro Leu Ser Arg Leu Glu Ile
290 295 300
Met Asp Arg Val Asn Arg Ser Arg Lys Phe Asp Thr Gln Phe His Gly
305 310 315 320
Phe Thr Gly Thr Asp Gly Pro Leu Gly Lys Arg Met Asp Asn Ser Lys
325 330 335
Thr Arg Ala Lys Leu Gly Trp Gln Pro Lys Tyr Pro Ser Phe Thr Glu
340 345 350
Phe Leu Gly Leu Ser Asn Leu
355
<210> 6
<211> 343
<212> PRT
<213> 葡萄
<400> 6
Met Gly Thr Leu Gly Ser Thr Thr Thr Leu Ser Pro Tyr Phe Ser Ser
1 5 10 15
Lys Leu Ser Phe Ser Ser Ser Phe His Arg Leu Arg Phe Ser Ala Ser
20 25 30
Arg Ser Phe Leu Ser Ile Phe Arg Asn Pro Ser Phe Arg Ala Lys Arg
35 40 45
Ser Val Ser Thr Asp Thr Arg Leu Arg Val Ser Ala Ser Ser Thr Leu
50 55 60
Gly Ala Pro Asn Glu Glu Met Glu Thr Ser Ser Phe Gly Leu Val Gly
65 70 75 80
Glu Asn Asp Leu Leu Ile Val Gly Pro Gly Val Leu Gly Arg Leu Val
85 90 95
Ala Glu Lys Trp Arg Glu Glu His Pro Gly Cys Gln Ile Tyr Gly Gln
100 105 110
Thr Met Thr Thr Asp His His Asp Glu Leu Val Lys Ile Gly Ile Asn
115 120 125
Pro Ser Leu Lys Gly Val Lys Thr Thr His Gln Phe Pro Tyr Val Ile
130 135 140
Phe Cys Ala Pro Pro Ser Arg Thr Ser Asp Tyr Pro Ala Asp Val Arg
145 150 155 160
Leu Ala Ala Ser Asn Trp Ser Gly Glu Gly Ser Phe Leu Phe Thr Ser
165 170 175
Ser Ser Ala Pro Phe Asp Cys Asn Asp Asn Gly Ser Cys Asp Glu Asp
180 185 190
Gly Pro Val Val Pro Ile Gly Arg Ser Pro Arg Thr Asp Val Leu Leu
195 200 205
Asn Ala Glu Lys Gly Val Leu Glu Phe Gly Gly Cys Val Leu Arg Leu
210 215 220
Ala Gly Leu Tyr Lys Ala Asp Arg Gly Ala His Val Tyr Trp Leu Lys
225 230 235 240
Lys Gly Thr Val Glu Ala Arg Pro Asp His Ile Leu Asn Leu Ile His
245 250 255
Tyr Glu Asp Ala Ala Ser Leu Ala Val Ala Ile Leu Lys Lys Lys Arg
260 265 270
His Gly Gln Ile Phe Leu Gly Cys Asp Asn His Pro Val Ser Arg Gln
275 280 285
Glu Leu Met Asp Leu Val Asn Lys Ser Gly Lys Phe Ser Lys Lys Phe
290 295 300
Glu Ala Phe Thr Gly Thr Ser Asp Pro Leu Gly Lys Arg Leu Asn Asn
305 310 315 320
Ser Lys Thr Arg Glu Glu Ile Gly Trp Gln Pro Lys Tyr Pro Ser Phe
325 330 335
Ser Gln Phe Leu Glu Ser Ile
340
<210> 7
<211> 304
<212> PRT
<213> 大叶钻天杨毛果杨亚种
<400> 7
Met Ala Ala Pro Leu Gln Val Ser Ala Phe Ser Thr Ile Gly Ala Arg
1 5 10 15
Asn Glu Glu Leu Gly Thr Ala Ser Ser Gly Leu Val Gly Glu Asn Asp
20 25 30
Leu Leu Ile Val Gly Pro Gly Val Leu Gly Arg Leu Val Ala Glu Lys
35 40 45
Trp Arg Gln Glu His Pro Gly Cys Gln Val Tyr Gly Gln Thr Val Thr
50 55 60
Thr Asp His His Asp Glu Leu Ile Lys Met Gly Ile Asn Pro Ser Leu
65 70 75 80
Lys Gly Thr Lys Ala Thr Gln Gln Tyr Pro Tyr Val Ile Phe Cys Ala
85 90 95
Pro Pro Ser Arg Thr Ser Asp Tyr Pro Gly Asp Val Arg Glu Ala Ala
100 105 110
Leu Ser Trp Asn Gly Asp Gly Ser Phe Val Phe Thr Ser Ser Ser Ala
115 120 125
Pro Tyr Asp Cys Phe Asp Asn Gly Gln Cys Asn Glu Asp Ser Pro Val
130 135 140
Val Pro Ile Gly Arg Ser Pro Arg Thr Asp Val Leu Leu Lys Ala Glu
145 150 155 160
Lys Val Val Leu Glu Ser Gly Gly Cys Ala Ile Arg Leu Ala Gly Leu
165 170 175
Tyr Ile Ser Phe Ser Val Leu Asn Tyr Val Asp Phe Ile Asn Asn Arg
180 185 190
Gly Ala His Ala Tyr Trp Leu Glu Lys Gly Thr Val Glu Val Arg Pro
195 200 205
Asp His Ile Leu Asn Leu Ile His Tyr Glu Asp Ala Ala Ser Leu Ala
210 215 220
Val Ala Ile Leu Lys Lys Lys Leu Arg Ser Arg Ile Phe Leu Gly Cys
225 230 235 240
Asp Asn His Pro Leu Ser Arg Gln Glu Val Met Asp Leu Val Ala Lys
245 250 255
Ser Gly Lys Phe Ser Lys Lys Phe Val Ala Phe Thr Gly Thr Ser Asp
260 265 270
Pro Leu Gly Lys Arg Leu Asn Asn Ser Lys Thr Arg Glu Glu Ile Gly
275 280 285
Trp Glu Pro Glu Tyr Pro Ser Phe Ala His Phe Leu Gly Val Ser Lys
290 295 300
<210> 8
<211> 351
<212> PRT
<213> 拟南芥
<400> 8
Met Gly Phe Ile Ser Cys Ile Ser Phe Pro Thr Ile Asn Ser Arg Ile
1 5 10 15
Leu Ser Thr His His Phe Ser Lys His Ser Thr Ser Ala Ser Ser Ser
20 25 30
Tyr Ser Leu Lys Phe Ala Leu Arg Arg Gln Glu Asp Lys Pro Lys Val
35 40 45
Ser Phe Phe Leu Pro Leu Thr Ser Ser Leu Met Ala Thr Pro Ile Gln
50 55 60
Ala Ser Ser Ser Ser Thr Ile Gly Glu Thr Ser Asp Gly Leu Lys Val
65 70 75 80
Gln Ser His Val Ser Ile Gly Ala Asn Asp Leu Leu Ile Val Gly Pro
85 90 95
Gly Val Leu Gly Arg Leu Val Ala Glu Gln Trp Arg Gln Glu His Pro
100 105 110
Glu Ser Gln Ile Phe Gly Gln Thr Val Thr Thr Asn His His Gly Glu
115 120 125
Leu Glu Asn Leu Gly Ile Lys Pro Ser Leu Lys Gly Thr Glu Tyr Gly
130 135 140
Gly Lys Phe Ser Tyr Val Ile Phe Cys Ala Pro Pro Ser Gln Ser Ala
145 150 155 160
Asp Tyr Ala Gly Glu Val Arg Asn Ala Ala Ser Asn Trp Asn Gly Glu
165 170 175
Gly Ser Phe Leu Phe Thr Ser Ser Ser Ala Pro Tyr Asp Cys Phe Asp
180 185 190
Asn Gly Glu Cys Asn Glu Asp Ser Pro Val Val Pro Leu Gly Lys Ser
195 200 205
Pro Arg Thr Asp Val Leu Leu Lys Ala Glu Lys Val Val Leu Glu Cys
210 215 220
Gly Gly Thr Val Leu Arg Leu Ala Gly Leu Tyr Thr Glu Thr Arg Gly
225 230 235 240
Ala His Thr Tyr Trp Leu Ser Lys Glu Thr Ile Asp Ala Arg Pro Asp
245 250 255
His Ile Leu Asn Leu Ile His Tyr Glu Asp Ala Ala Ser Leu Ala Val
260 265 270
Ala Ile Met Lys Lys Lys Ala Gly Ala Arg Ile Phe Leu Gly Cys Asp
275 280 285
Asn His Pro Leu Ser Arg Gln Glu Val Met Asp Leu Met Ala Gln Ser
290 295 300
Gly Lys Phe Asp Lys Lys Phe Lys Gly Phe Thr Ser Thr Ser Gly Pro
305 310 315 320
Leu Gly Lys Lys Leu Asn Asn Ser Lys Thr Arg Ala Glu Ile Gly Trp
325 330 335
Glu Pro Lys Tyr Pro Ser Phe Ala Gln Phe Phe Gly Val Ser Thr
340 345 350
<210> 9
<211> 287
<212> PRT
<213> 蓖麻
<400> 9
Met Ala Asn Pro Phe Gln Val Ser Ala Ser Ser Thr Met Gly Ala Thr
1 5 10 15
Asn Glu Glu Leu Asp Ala Val Ser Ser Ser Leu Val Gly Glu Asn Asp
20 25 30
Leu Leu Ile Val Gly Pro Gly Val Leu Gly Arg Leu Val Ala Glu Lys
35 40 45
Trp Arg Gln Glu His Pro Gly Cys Gln Val Tyr Gly Gln Thr Leu Thr
50 55 60
Thr Asp His His Asp Glu Leu Ile Lys Ile Gly Ile Asn Pro Ser Leu
65 70 75 80
Lys Gly Thr Lys Pro Ile His Gln Phe Pro Tyr Val Ile Phe Cys Ala
85 90 95
Pro Pro Ser Arg Thr Ser Asp Tyr Pro Gly Asp Val Arg Glu Ala Ala
100 105 110
Leu Ser Trp Asn Gly Glu Gly Ser Phe Leu Phe Thr Ser Ser Ser Ala
115 120 125
Pro Tyr Asp Cys Tyr Asp Asn Gly Asp Cys Asp Glu Asp Ser Pro Val
130 135 140
Val Pro Ile Gly Arg Ser Pro Arg Thr Asp Val Leu Leu Lys Ala Glu
145 150 155 160
Lys Val Val Leu Glu Ser Asp Gly Cys Val Tyr Lys Ala Asp Arg Gly
165 170 175
Ala His Val Tyr Trp Leu Gln Lys Gly Ile Val Glu Val Arg Pro Asp
180 185 190
His Ile Leu Asn Leu Ile His Tyr Glu Asp Ala Ala Ser Leu Ser Ile
195 200 205
Ala Ile Leu Lys Lys Lys Phe His Gly Arg Ile Phe Leu Gly Cys Asp
210 215 220
Asn His Pro Leu Ser Arg Gln Glu Val Met Asp Leu Val Ala Lys Ser
225 230 235 240
Gly Lys Phe Ser Lys Lys Phe Glu Ala Phe Thr Gly Thr Gly Asp Pro
245 250 255
Ser Gly Lys Arg Leu Asn Asn Ser Lys Thr Arg Glu Glu Val Gly Trp
260 265 270
Glu Pro Asn Tyr Pro Ser Phe Ala His Phe Leu Gly Val Ser Asp
275 280 285
<210> 10
<211> 284
<212> PRT
<213> 小立碗藓
<400> 10
Met Ser Gly Lys Glu Thr Glu Leu Glu Cys Asn Glu Glu Arg Ser Gln
1 5 10 15
Ile Thr Thr Asn Ser His Ala Asp Leu Leu Val Val Gly Pro Gly Val
20 25 30
Leu Gly Ser Leu Val Gly Arg Arg Trp Leu Glu Leu His Glu Gly Cys
35 40 45
Arg Val Val Gly Gln Thr Asn Thr Thr Asn Arg His Glu Glu Leu Leu
50 55 60
Ser Leu Gly Ile Phe Pro Val Thr Lys Asp Ser His Ser Gly Asp Lys
65 70 75 80
Phe Pro Tyr Val Ile Phe Cys Ala Pro Pro Ser Gly Ser Glu Asn Tyr
85 90 95
Ala Ala Glu Val Arg Ala Ala Ala Gln Arg Trp Asn Gly Glu Gly Ser
100 105 110
Leu Leu Phe Thr Ser Ser Ser Phe Val Tyr Asp Val His Asp Asn Gly
115 120 125
His Cys Asp Glu Ser Ala Pro Ile Thr Glu Lys Gly Thr Ser Pro Arg
130 135 140
Gly Asp Arg Leu Leu Asn Ala Glu Glu Glu Val Leu Lys Val Asp Gly
145 150 155 160
Asn Val Val Arg Leu Ala Gly Leu Tyr Ala Arg Asp Arg Gly Ala His
165 170 175
Met Tyr Trp Leu Gln Lys Gly Thr Val Asp Ala Arg Pro Asp His Phe
180 185 190
Leu Asn Leu Ile His Tyr Glu Asp Ser Ala Asp Leu Cys Ile Glu Ile
195 200 205
Leu Arg Lys Asn Leu Arg Gly Gln Ile Phe Met Gly Cys Asp Asn Thr
210 215 220
Pro Val Ser Arg Gln Asp Ile Met Asp Ile Met Met His Ser Gly Lys
225 230 235 240
Phe Ala Gly Asn Phe His Gly Phe Thr Lys Ser Asp Gly Pro Leu Gly
245 250 255
Lys Lys Met Asn Asn Ser Gln Thr Arg Glu Arg Leu Gly Trp Gln Pro
260 265 270
Lys Tyr Asn Ser Phe Lys Asp Tyr Val Ser Thr Leu
275 280
<210> 11
<211> 355
<212> PRT
<213> Ostreococcus tauri
<400> 11
Met Ser His Ala Leu Met Ser Ala Arg Ala His Val Thr Asn Val Ile
1 5 10 15
Gln Thr Ser Ser Arg Ile Arg Lys Arg Ser Cys Thr Ser Lys His Phe
20 25 30
Pro Ser Phe Arg Ala Ala Ala Ala Ala Ser Thr Ser Ser Ala Val Cys
35 40 45
Ala Ser Phe Thr Pro Pro Gly Ala Arg Glu His Asp Arg Asp Leu Leu
50 55 60
Ile Val Gly Pro Gly Val Leu Gly Ser Arg Ile Ala Arg Val Trp Leu
65 70 75 80
Glu Lys Tyr Pro Gly Ala Val Val Val Gly Gln Thr Asn Thr Thr Asn
85 90 95
Ala His Ala Gly Leu Thr Ser Ile Gly Val Ser Pro Arg Thr Lys Asp
100 105 110
Phe Asp Asp Asp Glu Pro Ser Ala Asn Arg Met Phe Pro Tyr Val Ile
115 120 125
Phe Ser Ala Pro Pro Ser Gly Ser Asp Asp Tyr Ala Gly Glu Val Glu
130 135 140
Ala Ala Leu Arg Tyr Trp Asn Gly Gly Gly Ala Phe Ala Phe Thr Ser
145 150 155 160
Ser Ser Ala Val Tyr Lys Asn Glu Ser Gly Asp Ala Cys Asp Glu Asp
165 170 175
Ser Glu Thr Tyr Asp Leu Gly Thr Asn Pro Arg Val Asp Arg Leu Leu
180 185 190
Lys Ala Glu Arg Ile Val Leu Asp Ala Gly Gly Val Val Cys Arg Leu
195 200 205
Ala Gly Leu Tyr His Ser Asp Arg Gly Ala His Lys Tyr Phe Ile Lys
210 215 220
Thr Pro Ser Ile Asp Ser Arg Ala Asp Ala Leu Val Asn Leu Ile His
225 230 235 240
Tyr Glu Asp Ala Ala Asp Leu Cys Val Ala Ala Met Asn Asn Gly Ser
245 250 255
Lys Ser Ala Val Tyr Leu Gly Thr Asp Gly Val Pro Ile Thr Arg Gly
260 265 270
Asp Ile Ala Arg Val Ala Val Glu Ser Gly Ala Tyr Gly Ala Asp Ala
275 280 285
Arg Ala Pro Ser Phe Thr Lys Thr Glu Gly Pro Ile Gly Arg Val Met
290 295 300
Ser Asn Asp Arg Thr Arg Thr Ala Leu Gly Trp Ala Pro Lys Tyr Val
305 310 315 320
Ser Phe Glu Thr Phe Met Thr Arg Val Asn Ala Arg Asp Ala Tyr Ser
325 330 335
Ala Ser Glu Lys Arg Pro Val Gly Trp Ala Pro Lys Gly Ser Ala His
340 345 350
Ile Ala Thr
355
<210> 12
<211> 344
<212> PRT
<213> Ostreococcus tauri
<400> 12
Met Arg Ala Ser Ser Ala Ser Pro Arg Ala His Arg Ala Phe Pro Ser
1 5 10 15
His Thr Ala Arg Lys Thr Ser Arg Glu Lys Ala Ser Ser Arg Ser Arg
20 25 30
Ala Val Ala Ala Ala Ser Thr Ser Asp Ala Pro Gly Pro Phe Gly Asp
35 40 45
Asp Arg Asn Leu Leu Val Val Gly Pro Gly Val Leu Gly Ser Arg Ile
50 55 60
Ala Arg Val Trp Leu Ser Asn Phe Pro Gly Ala Val Val Val Gly Gln
65 70 75 80
Thr Asn Thr Asp Ala Ala His Asp Gly Leu Arg Ser Val Gly Val Thr
85 90 95
Pro Arg Thr Lys Asp Phe Gly Ala Asp Asp Pro Thr Ala Thr Arg Arg
100 105 110
Phe Pro Tyr Val Val Phe Ser Ala Pro Pro Ser Gly Ser Glu Asp Tyr
115 120 125
Pro Gly Glu Val Ala Ala Ala Leu Lys Tyr Trp Asp Gly Ser Gly Ala
130 135 140
Phe Ala Phe Thr Ser Ser Ser Ala Val Tyr Lys Asn Glu Ala Gly Glu
145 150 155 160
Ala Cys Asp Glu Glu Ser Glu Val Tyr Glu Ile Gly Thr Asn Pro Arg
165 170 175
Val Asp Arg Leu Leu Lys Ala Glu Lys Val Val Leu Asp Ala Gly Gly
180 185 190
Val Val Cys Arg Leu Ala Gly Leu Tyr His Ser Glu Arg Gly Ala His
195 200 205
Lys Tyr Phe Ile Lys Thr Ser Ser Leu Asp Ser Arg Ala Asp Ala Leu
210 215 220
Val Asn Leu Ile His Tyr Glu Asp Ala Ala Asp Leu Cys Phe Ala Ala
225 230 235 240
Met Thr Lys Gly Ala Lys Ser His Ile Tyr Leu Gly Thr Asp Gly Val
245 250 255
Pro Ile Thr Arg Glu Ala Ile Ala Arg Val Ser Val Glu Ser Gly Val
260 265 270
Tyr Gly Ala Asp Ala Ala Ala Pro Ala Phe Thr Lys Thr Asp Gly Pro
275 280 285
Leu Gly Arg Ala Met Ser Asn Ser Arg Thr Lys Thr Glu Leu Asp Trp
290 295 300
Ser Pro Arg Tyr Glu Ser Phe Glu Ser Phe Ala Leu Arg Gln Gly Ala
305 310 315 320
Arg Asp Ser Tyr Ala Pro Trp Asn Ala Pro Thr Arg Ser Arg Gly Trp
325 330 335
Thr Pro Ala Gly Ala Arg His Val
340
<210> 13
<211> 294
<212> PRT
<213> Micromonas pusilla
<400> 13
Met Ala Ala Val Ser Glu Ser Cys Cys Arg Asp Leu Leu Val Val Gly
1 5 10 15
Pro Gly Val Leu Gly Ser Leu Val Cys Gln Arg Trp Leu Lys Thr Phe
20 25 30
Pro Ala Ala Thr Val Ile Gly Gln Thr Asn Thr Asp Ala Ser His Glu
35 40 45
Arg Leu Val Ala Leu Gly Ile Ser Pro Arg Leu Lys Ala Asp Ala Gly
50 55 60
Glu Ser Arg Arg Phe Pro Phe Val Val Phe Ser Ala Pro Pro Ser Gly
65 70 75 80
Ser Asp Asp Tyr Thr Ala Glu Val Glu Ala Ala Leu Lys Leu Trp Asp
85 90 95
Gly Thr Gly Gly Phe Val Phe Thr Ser Ser Thr Ala Val Tyr Ala Gly
100 105 110
Lys Asp Gly Glu Asp Cys Asp Glu Thr Thr Ala Gln Phe Gln Ile Gly
115 120 125
Glu Ser Pro Arg Ala Asp Lys Leu Leu Asn Ala Glu Ala Ala Val Leu
130 135 140
Gly Ala Gly Gly Cys Val Val Arg Leu Ser Gly Leu Tyr His Ser Gln
145 150 155 160
Arg Gly Ala His Met Tyr Phe Leu Lys Thr Pro Thr Leu Ala Ser Arg
165 170 175
Pro Asp Ala Leu Val Asn Leu Val His Tyr Glu Asp Ala Ala Ala Ala
180 185 190
Cys Val Arg Ala Leu Ser Ala Gln Leu Glu Gly Ser Ser Glu Gly Gly
195 200 205
Glu Ile Phe Leu Ala Thr Asp Gly Val Pro Val Thr Arg Glu Lys Met
210 215 220
Val Glu Ala Cys Leu Ala Cys Pro Asp Ala Tyr Asp Asp Gly Ala Met
225 230 235 240
Pro Glu Phe Ser Val Ser Asp Gly Pro Leu Gly Lys Ser Met Thr Asn
245 250 255
Pro Gln Thr Arg Glu Lys Leu Gly Trp Glu Pro Val Tyr Pro Ser Phe
260 265 270
Val Glu Phe Val Ala Ala Gly Ala Lys Asp Ser Phe Tyr Pro Pro Lys
275 280 285
Lys Lys Asn Thr Trp Ser
290
<210> 14
<211> 293
<212> PRT
<213> Micromonas pusilla
<400> 14
Met Ser Ser Cys Thr Phe Ala Thr Pro Arg Val Glu Val Ile Arg Ser
1 5 10 15
Arg Gly Ser Pro Leu Ser Ala Arg Ala Ala Arg Ser Ser Ser Ser Ser
20 25 30
Lys Phe Pro Ala Ala Ser Val Ile Gly Gln Thr Asn Thr Asp Thr Ser
35 40 45
His Glu Arg Leu Leu Ser Leu Gly Val Phe Pro Arg Leu Lys Glu Lys
50 55 60
Ala Gly Asp Glu Gln Tyr Pro Phe Val Val Phe Ser Ala Pro Pro Ser
65 70 75 80
Gly Ser Glu Asp Tyr Ala Ala Glu Val Glu Ala Ala Leu Lys Tyr Trp
85 90 95
Asp Gly Ser Gly Ala Phe Val Phe Thr Ser Ser Thr Ala Val Tyr Ala
100 105 110
Gly Lys Asp Gly Glu Pro Cys Asp Glu Ser Thr Pro Gln Phe Glu Ile
115 120 125
Gly Glu Ser Pro Arg Ala Asp Arg Leu Leu Lys Ala Glu Ala Ala Val
130 135 140
Leu Ala Ala Gly Gly Ser Val Val Arg Leu Ala Gly Leu Tyr His Ser
145 150 155 160
Gln Arg Gly Ala His Met Tyr Phe Leu Lys Thr Pro Ser Leu Ala Ser
165 170 175
Asn Ala Asp Gly Leu Val Asn Leu Ile His Tyr Glu Asp Ala Ala Ala
180 185 190
Ala Cys Val Asp Val Leu Val Ala Gln Phe Glu Gly Arg Thr Gly Gly
195 200 205
Gly Glu Val Phe Leu Ala Thr Asp Gly Val Pro Val Thr Arg Lys Glu
210 215 220
Met Val Glu Cys Cys Leu Glu Ser Asp Ala Tyr Asp Gly Asn Met Pro
225 230 235 240
Glu Phe Thr Glu Asp Asn Gly Pro Leu Gly Lys Ser Met Asn Asn Pro
245 250 255
Gln Thr Arg Glu Lys Leu Gly Trp Val Pro Val His Ala Ser Phe Val
260 265 270
Glu Phe Val Glu Ala Gly Ala Thr Asp Ser Phe Tyr Pro Lys Arg Arg
275 280 285
Lys Ser Thr Trp Lys
290
<210> 15
<211> 380
<212> PRT
<213> 莱茵衣藻
<400> 15
Met Ser Pro Arg Ser Cys Leu Ser Ala Ser Pro Thr Ser Ser Val Ala
1 5 10 15
Thr Arg Thr Thr Phe Thr Ser Thr Cys Ile Pro Arg Pro Arg Ala Ala
20 25 30
Gly Val Gln Val Ser Ala Gln Leu Asn Ile Ser Arg Arg Ser Ala Ser
35 40 45
Ala Ala Ala Ile Ala Ser Val Ala Pro Leu Gly Met Thr Phe Pro Gly
50 55 60
Ser Ile Asp Gly Gly Ala Ala Arg Gly Ser Val Ala Ala Ala Ala Thr
65 70 75 80
Ser Ser Leu Ala Gly Ala Val Ala Gly Ser Pro Ser Asn Leu Asp Leu
85 90 95
Leu Val Val Gly Pro Gly Val Leu Gly Ser Val Leu Gly Arg Asp Trp
100 105 110
Leu Ala Ser Val Gln Gly Gly Thr Ala Thr Gly Leu Thr Asn Thr Asp
115 120 125
Arg Ser His Glu Arg Leu Arg Ala Met Gly Leu Thr Pro Ala Thr Arg
130 135 140
Ser Thr Leu Pro Pro Asn Lys Lys Tyr Ser Phe Val Ala Phe Ala Ala
145 150 155 160
Pro Pro Ser Gly Ser Glu Asp Tyr Val Ala Asp Ile Lys Ser Ala Leu
165 170 175
Ala Leu Trp Asp Gly Ser Gly Ser Phe Ile Phe Thr Ser Ser Met Ser
180 185 190
Val Cys Ala Val Asp Asp Gly Gly Ser Ala Thr Asp Glu His Cys Pro
195 200 205
Leu Val Pro Val Gly Ala Gly Pro Ser Thr Asp Lys Leu Arg Gly Ala
210 215 220
Glu Glu Ala Val Leu Ala Ala Gly Gly Asn Val Leu Arg Leu Val Gly
225 230 235 240
Leu Tyr His Lys Phe Arg Gly Ala His Thr Phe Phe Ile Lys Gln Gly
245 250 255
Thr Val Ala Arg Pro Gly Gly Tyr Val Val Asn Leu Leu His Tyr Glu
260 265 270
Asp Ala Ala Ala Leu Ala Ala Ala Ile Leu Arg Gly Asp Gly Ser Gly
275 280 285
Pro Phe Arg Gly Arg Ala Phe Leu Gly Thr Asp Gly His Pro Val Thr
290 295 300
Phe Glu Asp Met Val Glu Tyr Cys Phe Ala Gly Gly Ala Phe Glu Arg
305 310 315 320
Val Pro Val Ser Phe Thr Gly Thr Phe Pro Asp Gly Gly Lys Thr Gly
325 330 335
Arg Gly Lys Arg Val Asp Asn Ser Gly Thr Ser Gln Ala Leu Gly Gly
340 345 350
Trp Lys Pro Lys Tyr Glu Ser Phe Gln Ser Phe Met Ala Ala Gly Gly
355 360 365
Ala Asp Tyr Tyr Asn Thr Ser Gly Leu Lys Trp Asn
370 375 380
<210> 16
<211> 292
<212> PRT
<213> 三角褐指藻
<400> 16
Met Ala Thr Lys Thr Ser Ser Gly Cys Ala Val Ile Gly Val Gly Val
1 5 10 15
Leu Gly Thr Ser Leu Cys Gln Gln Ile Leu Ser Ala Pro Glu Phe Asp
20 25 30
Gly Ile Lys Leu Thr Gly Ile Thr Lys Thr Thr Thr Asn His Asn Ala
35 40 45
Ile Arg Glu Lys Val Gly Met Asp Ser Glu Asp Arg Phe Gln Leu Leu
50 55 60
Thr Thr Asp Glu Cys Glu Gly Thr Glu Thr Lys Phe Lys His Ile Val
65 70 75 80
Phe Cys Ala Pro Pro Ser Gly Ser Glu Asp Tyr Pro Ala Asp Val Arg
85 90 95
Lys Ser Ala Asp Thr Leu Trp Ala Gly Pro Glu Glu Gly Gly Val Phe
100 105 110
Val Phe Thr Ser Ser Gly Ala Val Tyr Gly Pro Gly Asp Ser Arg Thr
115 120 125
Val Ser Glu Thr Ser Asp Ile Ala Asp Pro Glu Ser Ser Val Arg Val
130 135 140
Gly Arg Leu Val Lys Ala Glu Lys Ala Ala Leu Asp Ala Gly Gly Cys
145 150 155 160
Val Leu Arg Leu Ala Gly Leu Tyr Asn Leu Asp Arg Gly Ala His Asn
165 170 175
Phe Trp Leu Thr Ser Gly Lys Pro Ile Ser Gly Leu Pro Glu Gly Ile
180 185 190
Ile Asn Leu Leu His Tyr Glu Asp Ala Ala Ser Ala Cys Leu Ser Ala
195 200 205
Leu Lys Ala Gly Ser Ser Val Cys Glu Gly Arg Ala Phe Ile Ile Ser
210 215 220
Asp Gly His Pro Leu Thr Arg Lys Gln Ile Cys Glu Ser Ala Leu Gln
225 230 235 240
Ala Lys Thr Tyr Lys Asp Cys Ala Met Pro Thr Phe Ala Ser Glu Asn
245 250 255
Leu Asn Gly Met Ala Leu Gly Lys Val Tyr Asp Gly Ser Ser Ser Asn
260 265 270
Lys Ala Leu Glu Trp Ser Pro Arg Phe Glu Ser Phe Asp Thr Phe Met
275 280 285
Asn Ser Met Ala
290
<210> 17
<211> 274
<212> PRT
<213> 假微型海链藻
<400> 17
Met Ala Val Leu Ser Leu Leu Thr Ala Leu Leu Val Leu Ser Pro Ala
1 5 10 15
Arg Ala Phe Ser Thr Pro Gln Pro Ile Thr Ser Asp Leu Ala Ile Val
20 25 30
Gly Cys Gly Val Leu Gly Thr Ser Leu Cys Lys Gln Leu Leu Ser His
35 40 45
Pro Asp Phe Ser Ser Arg Ser Ile Thr Ala Ile Thr Lys Thr Thr Gly
50 55 60
Arg His Asp Ala Ile Arg Ala Glu Val Gly Asp Gly Asp Asp Thr Asp
65 70 75 80
Arg Phe Ala Val Leu Thr Met Asp Asp Val Leu Ala Gln Tyr Ser Gly
85 90 95
Asn Ser Phe Lys Asp Val Val Phe Cys Ala Pro Pro Ser Gly Phe Asp
100 105 110
Asp Tyr Pro Gln Ala Val Lys Asp Ala Ala Thr Gln Leu Trp Ser Gly
115 120 125
Pro Ser Ser Gly Gly Ser Phe Val Phe Thr Ser Ser Gly Gly Val Tyr
130 135 140
Glu Gly Leu Asp Gly Glu Thr Val Asn Glu Ser Ser Pro Thr Leu Asp
145 150 155 160
Ala Glu Ala Asn Pro Arg Gln Gly Arg Leu Ile Asn Ala Glu Arg Glu
165 170 175
Cys Ile Ala Leu Gly Gly Cys Ala Leu Arg Leu Ala Gly Leu Tyr Thr
180 185 190
Leu Glu Arg Gly Ala His Asn Tyr Trp Leu Glu Lys Cys Thr Glu Gly
195 200 205
Val Gln Gly Arg Glu Asp Gly Ile Val Asn Leu Leu His Tyr Asp Asp
210 215 220
Ala Ala Ser Ala Cys Leu Ala Ala Leu Gln Val Gly Pro Asp Val Asn
225 230 235 240
Ser Lys Gln Thr Tyr Leu Ile Ser Asp Gly Asn Pro Thr Thr Arg Lys
245 250 255
Gly Ile Cys Glu Ser Ala Leu Lys Ser Ala Arg Trp Phe Glu Gly Glu
260 265 270
Asn Leu
<210> 18
<211> 1397
<212> DNA
<213> 水稻
<400> 18
ctgctccaat ccactctcct ccgtttgcct gatccgatcc cctcacctcc gtccctgctt 60
cctcccggcg accaccgccc gccgcgcgat gggcggcgcc gccgtctcca gcctgcttgc 120
caccccaaca ccgacctctc gacctcgacc cgtctccacc accaccgccc ccttctccgt 180
caacctctcc accgcagctg cccgcgcacc tcgcctcctc ctcctctcgc gccgccctcg 240
ccctcgcccc gccgccgcgg ttctcggggt gtctgatgat acaggggtca agatggctgg 300
ctccgacatt gttggcaaga acgatttgct gattgttggc cctggagtgc ttggtcgact 360
ggtagctgag aaatggcagg aggaacatcc aggatgcaaa gtttttggcc agaccgcaag 420
cacagatcac cacaacgaat tgtcgaatat tggcatcatt ccctccttga agggatccac 480
ttttcctcag aaggttccat atgttatttt ctgtgctccc ccatctcgtt cggatgatta 540
ccctggggat gtgagagtag ctgcctcaaa ttggactggt gaaggctctt tcgtttttac 600
atcaagtact gctctgtacg attgtagtga caacgaattg tgcaatgagg attgcccatc 660
tgtgccaatt ggcagaagcc ctcgtactga cgtccttcta aaagcagaga atgttgttct 720
tgaggcagga ggctgtgtcc tcaggctagc aggactctat aaaatagata gaggtgctca 780
ttttttttgg ttgaggaaag gaactttgga cacacgacca gatcatatta tcaatcaaat 840
tcattatgag gatgctgctt cccttgcaat tgccataatg aaaaagggac acaggggtcg 900
aatctttttg ggctgtgaca ataagcctct ttccaggcaa gaaataatgg actctgttaa 960
cagaagtgga aaatttgaca cgaagttcca aggttttact ggtacagatg gtccactggg 1020
taagaagatg gagaattcga gaactcgttc tgagattggt tgggagccca agtatccaag 1080
cttcacagaa ttccttggtc ttgacagttg acatgcttgt gtatgcttgc aaaaatcttc 1140
aactctaaat gaacagcaac acgttgtgaa ccagttattt tgttttgttt ccattgcgtt 1200
ttttttaaaa gacgtaaata ctgtatatct tgttctgcta tgtgcttata tcagttagcg 1260
ttctaagtct gaccgttgta caatggtttc ggccagagaa atatttccac ccgtgtcatg 1320
ttggcctcgt tctggtccgt ccaacatcta tttgtctctt tgtaaccaca tttgttatga 1380
aaacttggtt tcctttc 1397
<210> 19
<211> 1425
<212> DNA
<213> 高梁
<400> 19
tactggaact cgatccactg tccatcttcc ccgtctactc tttcctcacc gtcacactgt 60
cgccgccgcc gccgcgaccg cgaccccgcc ggccgacgag atgagagccg ccgctgcggc 120
ctccttccat ctggcacccg ctacgaaccc ggcccatccc cgcggctcca cgacagactc 180
ctgttccctg aaacctgcac cgactgctca gccacctcgc ctccgttcgc tcgcccgccg 240
cgcgcctctc gtctgcgcct ctctcgggat atctcatgat aaagggtctg atatttccga 300
ccccaatgtt gttggccaga atgatttact gattgtgggc cctggtgttc ttggacgaat 360
tgtagctgag aagtggcaaa aggagcatcc aggttgcaaa gtttatggcc agactgcaag 420
caagaatcat cacagtgagt taacagatct tggcatcatc ccctcattga aaggcaccac 480
tattcatcag aaggttccac atgttatttt ctgcgctccc ccgtccagtt cagatgatta 540
tcctggggat gttagattgg cagcatcaaa ttggagtggt gaaggatctt tcctgtttac 600
atcaagtaca gctctgtatg actgcagtga caacagcatg tgcaacgagg attgttcgtc 660
agtgccaatt ggcaggagcc ctcgtactga tgtacttcta aaagtggaaa atgttgttct 720
tgaggcagga ggctgtgttc ttaggctagc tggactttat aagatagaca gaggtgctca 780
tgttttctgg ttgaggaaag gaacgttaga cacaagacca gatcatatca tcaatcagat 840
tcattatgag gatgccgctt cccttgcagt agcaatcatg aaaaagagac tgcggagtcg 900
gatatttttg ggctgtgaca acaagcccct ttccaggcaa gagataatgg atgctgttaa 960
caaaagtggg aaatttgaca cgaagtttga aggctttact ggtactgacg gtccattggg 1020
gaagagaatg gagaattcga aaactcgggc tgagatcggt tgggaaccga agtatccaag 1080
cttcaccgaa ttccttggta tcagcagtta acatccttgc atacttgtca gttactctga 1140
ctcagctgtg tagcaaccac gattagattg taccatcgtt tgaatttgat aatggtctca 1200
actggtgagg gtgcatatct tagtctactc cgggcctgtc ctcagtattt tttgattctc 1260
acactttttg tgttgtgaac cagactgatg caaccccacc cctccaggca acatatatat 1320
actgtcacgt gagtcgattt gtcgatcatg aacatatact ctgccgacta ccgtggtcgt 1380
ataactgcaa actagtttta aggtctggtc ctccggccag caagt 1425
<210> 20
<211> 994
<212> DNA
<213> 高梁
<400> 20
atgtccgccg ccgcagccga ctcctcctcc tcttcctcca tcggaatacc gcgtgatgcg 60
gcggttagct cgctctcgcc cgagagcgtc agcaacaacg atctgctcat cgttgggccg 120
ggcgtgctcg gccggatcgt cgccgagatg tggaaacagg aatatccagg ttgcaaggtt 180
tgtggccaga ctgcaaccac agatcatcac agtgaattaa ctgacattgg tatcattccc 240
tccttgaaga ggtccgtagc gggtcccaaa tttccaaatg ttattttctg tgctccacca 300
tatcgttctg aggattatgc tggagatctg agaatagcag cttcgaattg gaatggagaa 360
ggttctttcc tattcacctc gagtactgct gtgtatgact gcagtgacaa tggattctgc 420
ggtgaggatt ctccttgtgt atcgattggt cagagccctc gtactgatgt gcttctaaaa 480
gctgaaaatg ttgtccttga ggcagggggc tgtgttctta ggctagcagg actttacaaa 540
tcagatcgag gtcctcatgt ctattggttg tcaaaaggaa ccttggatgt gcgtcctgat 600
catatactca atctgataca ttatgaagat gcagcctctc ttgcaattgc catcatgaaa 660
aagagattac ggagccgcat ctttgtgggc tgtgacaatg agcctctgtc caggctagag 720
attatggacc gtgtcaacag aagcgggaaa tttgagacac agtttcaggg cttcactggg 780
actgatggtc cgctggggaa gaggatggag aactccaaaa ctcgggcaga gctcggatgg 840
cagcccaagt atccgagctt tacagagttc cttggtctca gcaatctcta actttcatgc 900
gttgtcgcta aatacttgtg attaaagatg aatatactgg tacatgaaag aaacaatgac 960
aaatttgaag tggaatgttg gttcctcata ctac 994
<210> 21
<211> 1275
<212> DNA
<213> 玉米
<400> 21
cacatcgccg ctgccgcgac ccgcgggccg acgagatgag agccgccgct gcggcctcct 60
tccatctggc acccgccacg aaaccggccc atcgccgcgg ctgcaccaca gagtcctgtt 120
ccctgaagcc tgcgccgact gctcggccac ctcgcctccg ctcgctcgcc ggccgggcgc 180
ctcttgtctg cgcctctctc gggatatctc atgataaagg gttcgacatt tctgacccca 240
atgttgttgg acagaatgat ttactgattg tgggccctgg tgttcttggg cgaatcatag 300
ctgagaagtg gaaaaaggag catccaagtt gcaaagttta tggccagacc gcaagcaaaa 360
atcatcacaa cgagttaaca gatcttggca tcatcccctc attgaaaggc accactgttc 420
atcagaaggt tccacatgtt attttctgcg ctcccccgtc tggttcagat gattacccta 480
gggatgtcag attggcagca tcaaattgga ctggtgaagg atctttcctg tttacatcaa 540
gtacagctct gtatgattgc agtgacaaca gcatgtgcaa cgaggattgt ctgtcagtgc 600
caattggcag gagccctcgt actgatattc ttctaaaagt ggaaaatgtt gttcttgagg 660
caggaggctg tgttctcagg ctggctggac tttataagat agacagaggt gcacatgttt 720
tctggttgag gaaaggaact ttagacacac gaccagatca tatcatcaac cagattcatt 780
atgaggatgc tgcctccctt gctgtagcaa tcatgaaaaa gggactgcgg agtcgaatat 840
ttctgggttg tgacaacaag cccctttcca ggcaagaaat aatggatgct gttaacaata 900
gtggaaaatt tgacacgaag tttggaggct ttactggtac tgacggtcca ttggggaaga 960
gaatggagaa ttcaaaaact cgggctgaga tcggttggga accaaagtat ccaagcttca 1020
ccgaattcct tggtatcagc agttaacatc cttgcatact gttcagttag tctgactcag 1080
ctgagtagca accgtgatta gattgtacca tcgtttgaat ttttatggcc tgaaccggtg 1140
atggtgtaca tctcagccta ctttgggcct gtccccagta tttttgattt tcacactatt 1200
tgagctatga accagacaga tgccacccca tcccatcgga taatatatat tgtcacgtga 1260
gtggatttgt cgatc 1275
<210> 22
<211> 1365
<212> DNA
<213> 玉米
<400> 22
ctccaacgtc cacgagacgc cgccacgcca tcggcatggg cagggcagca gcaccacttc 60
taccgcgtcc cattattgcc cagttccatc tccatcccca ccatctcgtc caccagtcgt 120
gtcttcccta tctccacgct accgcacctg tcgccgccgc ctccgccctc tttgctcccg 180
gccccacttc aagcccaagc gtccgcgtct cccgcccgcg ccgccacgcc tcgatgtccg 240
ccgccgccga ttcctcctcc tccattgtag tatcgggtga tgcagcggtt agctcgctct 300
cgcccgagag catcgagcac aacgatctgc tcatagttgg gcctggcgtg ctcggccgaa 360
tcgtcgctga gatgtggaaa caggaatatc caggttgcaa ggtttatggc cagactgcaa 420
ccacagatca tcacagtgaa ttgactgata ttggtatcat tccctccttg aaggggtccg 480
taccgggtcc aaaatttcca tatgttatct tctgtgctcc tccatatcgt tctgaggatt 540
atgctggaga tctgagagta gcagcttcaa attggaatgg aaaaggctct ttcctattca 600
cctcgagtac tgctgtgtat gactgcagtg acaatggatt ctgcagtgag gattctcctt 660
gtgtacccat tggtcaaagc actcgtactg atgtgcttct aaaagctgaa aatgttgtcc 720
ttgaggcagg cggctgtgtt cttaggctag taggacttta taaatcagat cggggtcctc 780
atgtttactg gctgtcaaaa ggaaccttgg atgtgcgtcc tgatcatata ctcaatctga 840
tacattatga agatgcagcc tctcttgtaa tttccatcat gaaaaagaga ttacggagct 900
gcatctttgt gggttgtgac aacgagcctc tgtccaggct agagatcatg gaccgtgtca 960
acagaagcag gaaatttgac acacagtttc atggcttcac tgggactgat ggtccgctgg 1020
ggaagaggat ggacaactcc aaaactcggg caaagcttgg atggcagccc aagtatccga 1080
gctttacaga gttccttggt ctcagcaatc tctaactttc atgcgatgtc actaaatacc 1140
tgttattaaa gatgaatata ctgatacata aaagagatga tgacagattt ggaagtggaa 1200
tgttgtttgc ttataccgca gttttcgact atcttttgat ttcttctgtt agatcttgta 1260
aagcattgca gcatggtgtg gcatgtacct gtgttgcctt ctattcgctt ctgttaactc 1320
atgaaagcat tgacttcaat acctgttaat tgtttccatg gcttc 1365
<210> 23
<211> 1294
<212> DNA
<213> 葡萄
<400> 23
atgggcactc ttggttcgac aaccactctc tctccgtact tctcctccaa gctctccttt 60
tcctcttcgt ttcatcggct ccgcttcagc gcctccagat cgttcctctc tatcttcaga 120
aaccctagct ttcgtgcgaa acgcagtgtt tcaacagaca ctcgtcttcg agtcagcgcg 180
tcgtccacgc ttggtgcacc aaacgaagaa atggagactt cttcttttgg tttggttgga 240
gagaatgacc ttctgattgt tggacctggt gttcttgggc gcttggtagc ggaaaaatgg 300
cgggaggaac atccaggatg tcaaatttat ggtcaaacta tgactacaga tcatcatgat 360
gaattggtta aaattgggat aaatccatct ttgaagggag tgaaaacaac tcatcagttt 420
ccatatgtca ttttctgtgc tccaccctcc cgcacctcag actaccctgc ggatgttagg 480
ttggctgcat caaactggag tggtgaaggt tctttcctat tcacatctag ttctgcacca 540
tttgattgca atgacaatgg atcatgtgat gaggatggtc ctgtagtgcc aattgggagg 600
agtcctagaa cagatgtcct tctaaatgca gaaaaaggag tactggagtt tggtggatgt 660
gttcttagat tggcaggact ttacaaagca gatagaggtg cacatgttta ttggttgaag 720
aaagggactg ttgaagcccg ccctgatcac atcctcaatc ttatacacta tgaggatgca 780
gcttctctag cagttgcaat tttgaagaag aaacggcatg gtcagatttt cttgggttgt 840
gataatcatc ccgtatccag gcaggaatta atggacttgg ttaataaaag cgggaaattc 900
agtaaaaagt ttgaggcttt tacaggaact agtgatcctt tgggtaagag attaaacaac 960
tcaaaaactc gtgaggaaat aggatggcag cctaaatacc caagcttctc gcagttcctt 1020
gagagtatct gagtaactgc tgcttctatt ttgtctttgg atttaatcca agtacggtgc 1080
aaaaagcaca tggattgctt gaagatggag ttgctgacct ttcttttctt ggcttacatg 1140
gcaaaatcct gctgcaacta aaaatttgtc aatagtcaac atgtaaatgt gatcacaagt 1200
tacctagtga aaaaatatct tttggcacac atataattgt cccatggaca tttgcttgag 1260
ttcataattc ttaccttgtc tactatacgc ttta 1294
<210> 24
<211> 1214
<212> DNA
<213> 毛果杨
<400> 24
ctaacgttat cttcaatggc agcacctctt caagtctcag cgttctctac tattggtgca 60
agaaatgagg agttggggac cgcaagttct ggtcttgttg gggagaatga tttgctgatt 120
gttggtcctg gtgttcttgg tcgcttagtc gctgagaaat ggcgccagga acatccgggt 180
tgtcaagttt atggccagac ggtgactaca gatcaccatg atgaattgat taaaatgggt 240
atcaatccat ctttgaaagg gactaaagca acacagcagt atccttatgt cattttctgt 300
gctccgcctt ctcgaacttc ggattaccct ggtgatgtaa gagaagctgc cttgagctgg 360
aatggggatg gttctttcgt gtttacatca agctctgcac cgtatgattg ttttgacaat 420
ggacagtgca atgaggactc tccggtagtg cccattggga gaagccccag aacagatgtc 480
cttctgaaag cagaaaaagt ggtgctggag agtggtggtt gtgctattag attggcagga 540
ctttatatat ccttctcagt ccttaattat gtggatttca taaacaatag aggtgcgcat 600
gcttactggt tggagaaggg tactgttgaa gttcgtccag atcacatcct gaatcttatt 660
cactatgagg atgctgcttc ccttgctgtt gcaatcttga agaagaaact tcggagccgg 720
attttcttgg gttgtgacaa ccatccatta tctaggcaag aagttatgga cttggtcgct 780
aaaagcggga agtttagcaa aaagtttgtg gcctttacag ggacaagtga tcctttaggc 840
aagagattaa acaactctaa gactcgcgag gagataggct gggagccaga ataccctagc 900
ttcgctcatt ttcttggtgt ctcaaaatag atgctatgcc tgtcagcagt aatgtgatgg 960
catagagctg acgggaagtc cacggttgtg acggaaagga tgcttggtgc tggggcagct 1020
gatttctacg tacctcacat ttgaaggcaa caaaccacct ccattttcac tatttttggt 1080
tagcagctgc aatgagaacc ggaaacctga acatgtattt gatactactc tttatgaata 1140
acaagtcttt agaaacgttt agattgtgtc tttgatctct tgtcttgcaa ataaaataca 1200
ctttcatttg attc 1214
<210> 25
<211> 1305
<212> DNA
<213> 拟南芥
<400> 25
tctttcccca cgccaactca atctttttcg tcgtcaaaag ccaaagcttt ttgctttttt 60
gtcttaatgg gtttcatctc ttgcatctca tttccgacga tcaattcaag aattctatcg 120
actcaccact tttccaagca ctcgacttca gcgtcttctt catattcact taagtttgct 180
ttgagacgtc aggaggataa acccaaagtc tcgttttttc ttccattaac gtcttcgtta 240
atggcgaccc ctattcaagc ctcttcttcc tccaccattg gtgagaccag tgatggcttg 300
aaggtccagt ctcatgtttc aattggagca aacgatctgc tgattgttgg accgggtgtt 360
cttggacgct tagttgcaga acagtggaga caggaacatc cagagtctca aatctttggg 420
cagacagtaa caacaaatca tcatggtgag ttggagaatt tgggtatcaa accatctctt 480
aaaggaaccg aatatggggg aaagttctcc tatgtgatct tttgtgctcc accatcacaa 540
agcgctgatt atgctggtga agtcaggaat gcagcatcaa actggaatgg cgaaggatca 600
ttcttattca catctagttc tgcaccttat gattgctttg ataacggaga atgcaacgag 660
gattctccag tagtgccact gggaaagagt ccaagaactg atgtgctttt gaaagctgaa 720
aaagtagtgt tggaatgtgg agggactgtc cttagactag cagggcttta cacagaaact 780
agaggtgcac atacttactg gttgagtaag gagacaattg atgctcgtcc tgatcatatt 840
ctaaatctca tacactatga ggatgcagca tcgctggcag ttgcaatcat gaagaagaaa 900
gccggtgctc ggattttctt gggttgtgac aaccatcctt tgtcaaggca agaggtgatg 960
gacctgatgg ctcaaagcgg aaaatttgat aagaagttca aaggttttac aagcaccagt 1020
ggtcctttag ggaagaagct gaacaactct aagacacgag cggagatagg atgggagccg 1080
aagtatccaa gctttgccca attttttgga gtatcgacat aatattttta cttggattga 1140
ttaagaatgt ctctagcgct gaagaatcca ataatgtgaa gcattattta tgtttggtac 1200
aaacaaactc atgtatcctc ttgagtgatc aaacagctga aattttcaga attcttcata 1260
aagctccatg gttcatatat ataaatcata ttttgttaat ttact 1305
<210> 26
<211> 864
<212> DNA
<213> 蓖麻
<400> 26
atggctaacc cttttcaagt ttctgcttct tctacaatgg gtgcaacgaa tgaggaattg 60
gatgctgttt cttctagtct tgttggagag aatgatcttt tgattgttgg tcctggtgtt 120
cttggccgtt tagtagctga gaaatggcgt caagaacatc ctggttgtca agtttatggg 180
cagacattga ctacagatca ccatgacgag ttgatcaaaa tagggatcaa tccatctttg 240
aagggaacta aaccaattca tcagtttcct tatgtcatat tctgtgctcc tccctcgcga 300
acatctgact accctggtga tgtcagggaa gctgctttaa gctggaatgg agaaggttct 360
ttcttattta catcgagctc tgcaccatat gactgctatg acaacggaga ttgtgatgag 420
gactctccag tagtgccaat tgggcgaagc ccgaggacag acgtgcttct aaaggcagaa 480
aaagtagtgc tggagagtga tggctgtgtc tacaaagcag atagaggtgc acatgtttat 540
tggttgcaga aggggattgt agaagttcgt cctgatcaca tcttaaatct tatacactat 600
gaggatgcag cttctctctc cattgcaatc ttgaagaaga aattccatgg gcgaattttt 660
ttgggttgtg ataatcatcc attatccagg caggaagtga tggacttggt tgctaaaagt 720
gggaaattta gtaaaaagtt tgaggccttt actggaaccg gtgatccttc aggcaagaga 780
ttgaacaact cgaaaactcg agaggaagta ggatgggagc caaattaccc tagctttgct 840
cattttcttg gggtgtctga ctaa 864
<210> 27
<211> 1225
<212> DNA
<213> 小立碗藓
<400> 27
agagcttggc gtagcgtcgt cggccatttt ggagtttctt accacgatct ctcactattt 60
gcagcggcgt cgcagctcgt tcaatcgatc gtgatttcgt tgccgagatt atcgtttgct 120
aggttgcgtt cagacgcgtc aggtgcggag cgaaaatgtc ggggaaggaa acggagttag 180
agtgtaacga agaaaggagt caaattacga ccaactccca tgctgatttg ctggttgttg 240
gacctggagt gcttggtagt cttgtaggcc gacgttggtt agagctgcat gaagggtgta 300
gagttgtggg acagacgaat accaccaacc gacatgaaga attgctgtcg ctcggcattt 360
ttccagtcac aaaggattct cattctggag acaaatttcc ttatgtgata ttctgcgctc 420
caccgtctgg tagtgagaat tacgcagcag aagtcagggc agcagctcag aggtggaatg 480
gggaggggtc attgttattc acatctagca gttttgttta tgacgttcat gacaatggcc 540
attgtgatga gagtgcgccc ataactgaga aaggaacgag ccctcgtggg gatagattgc 600
tgaatgcgga ggaggaggtt ttgaaggtcg atggaaacgt tgtacgattg gcaggcctat 660
atgctcggga tcgtggagcg cacatgtact ggcttcaaaa aggaacagta gatgccaggc 720
ccgaccattt tttgaatctt attcattatg aggactcggc tgatttgtgc atcgaaattt 780
tgcggaaaaa tctacgtggt caaatcttca tgggttgtga taacacacct gtttctaggc 840
aggacattat ggacatcatg atgcacagtg gtaaatttgc aggaaatttc catggattca 900
ctaaaagtga tggtcctctc gggaaaaaga tgaataacag tcaaactagg gaaagacttg 960
ggtggcaacc aaagtacaat agcttcaagg attacgttag caccttgtga ggatgcggtc 1020
tgttatctca ttgtcgtcga ctttcattct gtagacattt taagacaatt ttctgcccta 1080
tttccgcatg gtgacttcgt taacctcatc acttggagaa tttttgtagc atcttagaca 1140
gtgaattgat ttatcttatt tcatgtgtca tgggtgaatg ttttaaccat gccatgtcca 1200
gtttgtggta aacgagcagt tattc 1225
<210> 28
<211> 1068
<212> DNA
<213> Ostreococcus tauri
<400> 28
atgtcgcacg ccttaatgag cgcccgcgcg cacgttacga acgtcatcca aacctcttcg 60
cgcattcgta agcgctcgtg cacttcaaag cactttccga gtttcagggc ggcggcggcg 120
gcgtcgacat cttccgccgt gtgcgcgtcg ttcacccctc cgggcgctcg cgagcacgac 180
cgggacttgc tcatcgtcgg tccaggggtt ctaggttcgc gaatcgcgag ggtatggttg 240
gagaagtacc caggggcggt cgttgtcggg cagacgaaca cgacgaacgc gcacgccggg 300
ttgacgtcca tcggggtgtc gcctcggacg aaggatttcg acgacgacga gccgagtgcg 360
aacaggatgt tcccgtacgt tattttcagc gcaccgccga gcgggagcga cgattacgcg 420
ggtgaggtag aggcggcgct gaggtattgg aacggcggag gggcgtttgc gtttacgagc 480
tcgagcgcgg tgtacaaaaa cgagagcggc gacgcgtgcg atgaggatag cgaaacgtac 540
gatttaggaa cgaacccgcg agtcgatcgc ttgctcaagg cagagcgcat cgttctcgac 600
gcgggcgggg ttgtttgccg attggccggt ctgtaccact cggaccgcgg ggcgcacaag 660
tacttcatca agacgccttc gatcgactct cgcgccgacg cgttggtgaa cttgatccac 720
tacgaggacg ccgccgatct gtgcgtcgcc gcgatgaaca acgggtcgaa gtctgcggtg 780
tacctcggta cggatggggt tccgatcacg cgaggcgata tcgctcgcgt cgccgtcgag 840
agcggcgcgt acggcgccga cgcgcgcgcg ccgtcgttca ccaaaaccga gggcccgatc 900
ggtcgcgtca tgtccaacga ccgaacgcga acggcgctcg gttgggcacc taaatacgtc 960
tctttcgaga cctttatgac gcgcgtcaac gctcgcgacg cgtactcggc gtcggagaag 1020
cgacccgtcg ggtgggcgcc aaagggcagc gcccacatcg cgacgtga 1068
<210> 29
<211> 1035
<212> DNA
<213> Ostreococcus tauri
<400> 29
atgcgcgctt cgagcgcgtc accgcgcgcg caccgcgcgt ttccctcgca caccgcgcga 60
aagacgtcgc gcgaaaaagc gtcgtctcgg tctcgcgcgg tggcggcggc gtcgacttcg 120
gacgcaccgg ggccgttcgg ggacgaccga aacctgctcg tcgtcggacc cggtgttttg 180
ggctcgcgca tcgcgcgcgt gtggctgtcg aacttccccg gggcggtggt ggtcggacag 240
acgaacaccg acgccgcgca cgacggcttg cgaagcgtcg gcgtgacgcc gcgaacgaag 300
gatttcggtg cggacgatcc cacggcgacg cggcggtttc cctacgtcgt tttcagcgcg 360
ccaccgagtg gcagtgagga ttacccgggc gaagtcgcgg cggcgttgaa gtattgggac 420
ggctcgggcg cgttcgcgtt tacgagctcg agcgccgtgt acaagaatga agccggcgag 480
gcgtgcgacg aggagagtga agtttatgaa ataggcacca atcctcgtgt cgatcggttg 540
ctgaaggcgg aaaaagtcgt gttagacgct ggtggtgtgg tgtgtcgatt agccgggctc 600
tatcactccg agcgaggcgc gcacaagtac ttcatcaaga cgtcctcgct cgattctcgc 660
gccgacgcct tggtgaattt aattcactac gaagacgccg ccgatctgtg cttcgccgcg 720
atgacgaaag gagcaaagtc tcacatttat ctcggcaccg acggcgtccc gatcacccgc 780
gaggcgatcg cgcgcgtctc cgtggagtct ggcgtctacg gcgccgacgc cgccgcgccc 840
gccttcacca aaaccgacgg cccgctcggt cgcgccatgt ccaactcccg caccaagacc 900
gagctcgatt ggtcgcctcg gtacgaatct ttcgaatctt tcgcgttgcg tcagggcgcg 960
cgcgactcgt acgcgccctg gaacgcgccg acgcgttcgc gcgggtggac ccccgcgggc 1020
gcgcgtcacg tgtga 1035
<210> 30
<211> 885
<212> DNA
<213> Micromonas pusilla
<400> 30
atggcggcgg tatccgagag ctgctgtcgc gatctcctcg tcgtaggccc cggagttctc 60
ggcagtctcg tgtgccaacg atggctcaag acgttccccg cggcgacggt gatcggccag 120
acgaacacgg acgcgtcgca cgagcggttg gtcgccctcg gaatctcgcc gcgtttgaaa 180
gcggacgcgg gggagtctcg ccgattcccg ttcgtcgtct tcagcgcgcc gccgtccgga 240
agcgacgact acaccgccga ggtcgaagcc gcgctgaagc tgtgggacgg caccggtgga 300
ttcgtgttca cgagcagcac cgcggtgtac gcgggtaaag acggcgagga ctgcgacgag 360
accacggcgc agttccaaat cggtgagtcc ccgagggcgg acaagttgct gaacgcggaa 420
gccgcggttc tgggcgccgg cgggtgcgtc gtgcgcctgt ccggtctgta tcactcccaa 480
cgcggggcac acatgtactt tttgaagacc ccgacgttgg cgtccagacc cgacgcgctg 540
gtgaacctcg tgcactacga agacgccgcg gccgcttgcg tgcgcgcgtt atccgcgcag 600
ctcgagggca gctcggaggg tggcgagata ttcctcgcca cggacggcgt gcccgttacg 660
cgagagaaga tggtcgaggc gtgtctggcg tgccccgacg cgtacgacga cggagcgatg 720
ccggagttta gcgtgagcga cgggccgctc gggaagagca tgacgaatcc gcaaacgcgc 780
gagaagctcg gctgggagcc ggtgtatccg agcttcgtcg agttcgtcgc tgcgggcgcg 840
aaagactcgt tctatcctcc gaagaagaag aacacgtgga gttag 885
<210> 31
<211> 882
<212> DNA
<213> Micromonas pusilla
<400> 31
atgtcgagct gcactttcgc cactccacgt gtcgaagtga ttaggtcgcg gggatctcct 60
ctcagcgcga gagccgcgcg ctcctcctcc tcgtccaaat ttccggcggc ttccgtgatc 120
ggtcagacaa ataccgacac ttcgcacgag aggctgctct ccctcggcgt gtttccccgg 180
ctgaaggaaa aggctgggga tgagcagtac cctttcgtcg tgttcagcgc gccgccttct 240
ggcagcgagg actacgccgc cgaggtcgag gcagccctga agtactggga tggcagcggc 300
gcttttgtct tcaccagcag cacggcggtg tatgccggca aggacggaga gccttgcgat 360
gagagcacgc ctcagtttga gattggggag tcacctcgcg cggataggct gctgaaggcc 420
gaggcggcgg tgctggccgc gggtgggagc gtcgttcgtc tcgcgggact gtatcactca 480
cagcggggtg ctcacatgta ctttctgaaa accccctctc ttgcatccaa cgctgacggt 540
ctggtcaacc tgattcacta cgaggacgcg gcggcggcgt gtgtcgacgt gctcgttgcg 600
cagtttgagg gaaggacagg cggaggggag gttttcctgg ccacagacgg cgtgcccgtt 660
accaggaagg aaatggtcga gtgctgcttg gagagcgatg cctacgacgg aaacatgccg 720
gaattcacgg aggacaatgg acccctgggt aagagcatga acaaccctca aactcgtgag 780
aaactcggct gggttccggt gcacgcgagc ttcgtcgagt ttgtcgaggc gggtgcgacg 840
gattcattct atcccaagcg caggaagagc acttggaagt ag 882
<210> 32
<211> 1578
<212> DNA
<213> 莱茵衣藻
<400> 32
agctgctcgt aatatctggg atcttgtgga gaggccagct cgggtcggtt ggagacgaca 60
tcatgtcgcc ccggagctgc ttgagcgctt cgcccaccag ctctgtcgct actcgcacta 120
cattcacatc tacctgtatt cctcgtcccc gagcggcggg agtccaagtc tcggcgcaac 180
tgaacatttc gcgtcgcagc gctagcgccg ctgctatcgc tagcgttgca ccgctgggga 240
tgacctttcc tggatcaatc gacggtggtg ccgcacgcgg ctctgtcgca gctgctgcaa 300
ccagctctct ggcgggagca gtggcggggt cgccgtccaa tctggaccta ctggtggtgg 360
ggccgggtgt gttgggcagc gtcctgggcc gcgactggct ggcgtctgtg cagggcggca 420
cagccaccgg cctcaccaac acggatcgca gccacgagcg gctgcgcgcc atggggctga 480
cgccggccac acgttccacc ctaccgccca acaagaaata cagcttcgtg gccttcgccg 540
cgccgccctc aggcagtgag gattacgttg ctgatatcaa gtcggctctg gcattgtggg 600
acggcagtgg ctcattcatc ttcacgtcgt ccatgtcggt gtgcgcagtg gacgacggtg 660
gctccgctac cgacgagcac tgcccgctgg tgcccgtggg cgcggggccc tccactgaca 720
agctgcgcgg ggcggaggag gctgtcctgg cggccggtgg caacgtgcta cgcctggtgg 780
gtctgtacca caagtttcgc ggcgcacaca ccttctttat caaacagggc actgtggccc 840
ggccaggggg ctatgtggtg aacctgctgc actacgagga cgctgcggca ctggctgcag 900
cgatcctgcg tggtgatggc tctgggccct tccgaggccg tgcgttcctg ggcactgatg 960
gccacccagt aacgtttgag gacatggtgg agtactgctt tgctggtggt gcctttgagc 1020
gcgtaccagt gtcgttcacc ggcaccttcc ctgacggtgg caagaccggt cggggcaagc 1080
gtgtagacaa cagcggaaca agccaggcgc taggtggctg gaagcccaag tacgagtcat 1140
tccagtcctt tatggctgct ggcggggctg actactacaa cacttcgggg ctaaagtgga 1200
actaaagtgg gagtgtaagc caggtgcaca cgatgggtga actgttagac ttgcgacaag 1260
cgtgtgggct tgaggcaatg ccacacgact gcagtggtgt tggcacaaga caggagcacg 1320
gaaatcgtgg gagtgcaggg ggcagacttg gtgtcagcag tagcagccac agaggttatg 1380
tcgaaggggc tgctagcgaa ctgaagatag agggcttcat tagttgcgca actgtggcac 1440
aggcaggcca ggcagaaggc atgactggtg ctagggtggc tgagagcacg ttgggaacaa 1500
ccgttccctg gcttcgcaac cttcccctgg cttgacgtgc agtcacgaag ggcactgtaa 1560
gcacgtttga gtgtgtga 1578
<210> 33
<211> 952
<212> DNA
<213> 三角褐指藻
<400> 33
tcttcgactc ataaaatcgt ccatcgtttc tgtatcgatt gcacttgctt ctattttagc 60
atcggctttg gccatggcga cgaagacatc atcggggtgt gctgtcatcg gagttggcgt 120
tctaggtacc agtttatgcc aacagatctt gtctgcgcct gagtttgatg gaatcaagct 180
tacaggcatt acaaaaacta ccaccaatca caacgcgatc cgagaaaaag tcggaatgga 240
tagcgaagat cgtttccagc tattgaccac agatgaatgt gaaggaacgg aaaccaaatt 300
caagcatatc gtgttttgtg cgccgccttc gggttctgaa gactacccag ccgacgtacg 360
gaaatccgcc gatacgctat gggcaggacc ggaagaaggg ggcgttttcg tgtttacctc 420
cagcggtgca gtatacggac ctggggattc tagaacagta tcagaaacat ccgatattgc 480
tgatcctgaa tcgagtgtac gagtcggacg cctcgtgaag gccgaaaagg ctgctctaga 540
cgctggcgga tgcgtgttgc ggttggctgg tctctacaat ttagatcgcg gcgctcacaa 600
tttttggttg accagcggga agccaatatc cgggctaccc gaaggcatca tcaatctact 660
gcattacgag gatgccgcaa gtgcctgtct atcggcgctc aaagctggct ccagcgtctg 720
cgaaggtcga gcctttataa tcagtgatgg tcatcctctc acacgaaaac aaatctgcga 780
aagcgcgctg caagcaaaaa cctataaaga ttgtgcaatg cctacatttg catccgaaaa 840
tttgaatggc atggccttgg ggaaggtgta cgatggatcc tcaagtaaca aggcgttaga 900
gtggtctccg cgctttgaat cctttgacac atttatgaac agcatggcct aa 952
<210> 34
<211> 825
<212> DNA
<213> 假微型海链藻
<400> 34
atggcggtgc tctccctcct aaccgctctc ctcgtgctct caccagcacg agctttctcc 60
acgccccaac ccatcacttc cgacctagcc attgtaggat gtggcgtcct cggcacatcg 120
ctatgcaaac aactgctatc acatcctgac ttttcatctc ggtccatcac ggccattacc 180
aagaccactg gtcgtcatga tgccattcgt gccgaggttg gagacggcga tgacacggat 240
agattcgcag tgctgacaat ggatgatgtg ttggctcaat acagtggaaa ttccttcaag 300
gatgttgtat tttgtgcacc gccatcgggt tttgacgact atccccaggc agtcaaagat 360
gcagcgacac agttgtggtc ggggccttcg tccggtgggt cgttcgtatt cacttccagt 420
ggtggagtgt atgaagggtt agatggggag actgtgaatg aatcatcgcc tacgttggat 480
gcagaggcaa atccaagaca ggggaggtta atcaacgcgg aacgtgaatg tattgcgttg 540
ggggggtgtg cactacgttt ggctgggctg tatactttgg aaagaggggc acacaactac 600
tggcttgaaa aatgtaccga gggagttcaa gggagagaag acggtattgt gaacctattg 660
cattacgatg acgctgcgtc ggcgtgtctc gccgcgttgc aggtggggcc tgatgtgaac 720
tccaaacaga cgtacttgat tagtgacggg aatccaacta cgcgaaaggg gatctgcgag 780
agtgccttga agagtgcaag atggtttgaa ggggaaaatt tatga 825
<210> 35
<211> 21
<212> DNA
<213> 合成的
<400> 35
gtcgtgccat gacatctacc a 21
<210> 36
<211> 21
<212> DNA
<213> 合成的
<400> 36
ctgctttatt gcctcacttg c 21
<210> 37
<211> 20
<212> DNA
<213> 合成的
<400> 37
cgtcgtgctt cctgactcca 20
<210> 38
<211> 21
<212> DNA
<213> 合成的
<400> 38
cctccaccaa catgctcctt c 21
<210> 39
<211> 24
<212> DNA
<213> 合成的
<400> 39
acataaccta ccgaagaaga gtgg 24
<210> 40
<211> 22
<212> DNA
<213> 合成的
<400> 40
tcacagcctg aagcacataa aa 22
<210> 41
<211> 21
<212> DNA
<213> 合成的
<400> 41
ggatacggca ctggatcttg g 21
<210> 42
<211> 21
<212> DNA
<213> 合成的
<400> 42
ttgaatggag ggtcgttgag c 21
<210> 43
<211> 22
<212> DNA
<213> 合成的
<400> 43
tactgctccg atacttcaac cc 22
<210> 44
<211> 22
<212> DNA
<213> 合成的
<400> 44
ccatccgcta gatcaacaac at 22
<210> 45
<211> 20
<212> DNA
<213> 合成的
<400> 45
ccaagacgcc ctggtgatgc 20
<210> 46
<211> 24
<212> DNA
<213> 合成的
<400> 46
ttcgctttcc agttgagttc cttc 24
<210> 47
<211> 21
<212> DNA
<213> 合成的
<400> 47
tcgctactcc tgacattggt t 21
<210> 48
<211> 20
<212> DNA
<213> 合成的
<400> 48
tgatcgccct aattctgctc 20
<210> 49
<211> 21
<212> DNA
<213> 合成的
<400> 49
tggaacagga aagggaacat c 21
<210> 50
<211> 21
<212> DNA
<213> 合成的
<400> 50
tcgtggacca ataaccaaag g 21
<210> 51
<211> 21
<212> DNA
<213> 合成的
<400> 51
gccagaacaa acccatcaaa c 21
<210> 52
<211> 21
<212> DNA
<213> 合成的
<400> 52
gtaactccag agccgaacca g 21
<210> 53
<211> 20
<212> DNA
<213> 合成的
<400> 53
ataatctccg atggctgttc 20
<210> 54
<211> 22
<212> DNA
<213> 合成的
<400> 54
tccagacctt atgtagtatc cc 22
<210> 55
<211> 19
<212> DNA
<213> 合成的
<400> 55
gggccttcat ggatcaacc 19
<210> 56
<211> 20
<212> DNA
<213> 合成的
<400> 56
ccgcttcaag catcctcatc 20
<210> 57
<211> 20
<212> DNA
<213> 合成的
<400> 57
gcctgccctg gactacattg 20
<210> 58
<211> 24
<212> DNA
<213> 合成的
<400> 58
gcaaacatat gtacacggtt ctgg 24
<210> 59
<211> 19
<212> DNA
<213> 合成的
<400> 59
aacgtggctg ctccttgaa 19
<210> 60
<211> 20
<212> DNA
<213> 合成的
<400> 60
ttggcaataa gccacacaca 20
<210> 61
<211> 19
<212> DNA
<213> 合成的
<400> 61
tggccatgga agagttggc 19
<210> 62
<211> 19
<212> DNA
<213> 合成的
<400> 62
cagaagcaac tgctccacc 19
<210> 63
<211> 20
<212> DNA
<213> 合成的
<400> 63
gaagtccttg agccgtcctg 20
<210> 64
<211> 20
<212> DNA
<213> 合成的
<400> 64
aagtcccttg atgccctcct 20
<210> 65
<211> 21
<212> DNA
<213> 合成的
<400> 65
agaagggatc cagatgaaga a 21
<210> 66
<211> 22
<212> DNA
<213> 合成的
<400> 66
aacaagaaac gagcaacata ga 22
<210> 67
<211> 20
<212> DNA
<213> 合成的
<400> 67
gcctccatca tcatcttcca 20
<210> 68
<211> 20
<212> DNA
<213> 合成的
<400> 68
attgttacat cccgagcacc 20
<210> 69
<211> 21
<212> DNA
<213> 合成的
<400> 69
gccagaacaa acccatcaaa c 21
<210> 70
<211> 21
<212> DNA
<213> 合成的
<400> 70
gtaactccag agccgaacca g 21
<210> 71
<211> 20
<212> DNA
<213> 合成的
<400> 71
gcttccacat tgacccatac 20
<210> 72
<211> 20
<212> DNA
<213> 合成的
<400> 72
cttgagggca tacagcatct 20
<210> 73
<211> 20
<212> DNA
<213> 合成的
<400> 73
cacgacgcag tgatctgagg 20
<210> 74
<211> 20
<212> DNA
<213> 合成的
<400> 74
gatgaaacgc agggaatacg 20
<210> 75
<211> 19
<212> DNA
<213> 合成的
<400> 75
gacatcgtca agagggtcg 19
<210> 76
<211> 20
<212> DNA
<213> 合成的
<400> 76
ccatccatga tccatcatcc 20
Claims (25)
1.一种分离的多核苷酸,其包含选自下述的核苷酸序列:
(a) 核苷酸序列,其编码具有UDP-葡萄糖差向异构酶活性的多肽,其中所述多肽具有与SEQ ID NO: 1相比至少70%序列同一性的氨基酸序列,这基于Clustal V比对方法,其中逐对比对的默认参数为:KTUPLE=1,GAP PENALTY=3,WINDOW=5和DIAGONALS SAVED=5;
(b) 核苷酸序列,其编码具有UDP-葡萄糖差向异构酶活性的多肽,其中所述核苷酸序列在严谨条件下可与包含SEQ ID NO: 18的完整互补体的DNA分子杂交;
(c) 核苷酸序列,其编码具有UDP-葡萄糖差向异构酶活性的多肽,其中通过至少一种选自缺失、置换、添加和插入的方法,通过改变一个或多个核苷酸,从SEQ ID NO: 18衍生出所述核苷酸序列;
(d) 编码多肽的核苷酸序列,其中所述多肽的氨基酸序列包含SEQ ID NO: 1;和
(e)包含SEQ ID NO: 18的核苷酸序列。
2.一种重组DNA构建体,其包含与至少一个调控元件可操作地连接的根据权利要求1所述的分离的多核苷酸。
3.一种增加植物产量的方法,所述方法包括:表达与至少一个调控元件可操作地连接的重组叶绿体UDP -葡萄糖差向异构酶,其中所述差向异构酶定位在叶绿体中。
4.根据权利要求3所述的方法,其中所述重组叶绿体UDP -葡萄糖差向异构酶包含叶绿体转运肽。
5.根据权利要求3所述的方法,其中所述叶绿体UDP -葡萄糖差向异构酶包含根据权利要求1所述的核苷酸序列。
6.一种在原位增加叶绿体中用于单半乳糖基二酰基甘油(MGDG)生物合成的UDP-半乳糖的生产的方法,所述方法包括:在至少一个调节启动子元件控制下,表达叶绿体UDP -葡萄糖差向异构酶,其中所述差向异构酶包含叶绿体转运肽。
7.根据权利要求6所述的方法,其中所述叶绿体UDP -葡萄糖差向异构酶包含根据权利要求1所述的核苷酸序列。
8.一种改变植物的农艺学特性的方法,所述方法包括:表达包含根据权利要求1所述的核苷酸序列的UDP-葡萄糖差向异构酶。
9.根据权利要求8所述的方法,其中所述农艺学特性选自:增加的光合效率、产量和干旱耐受性。
10.一种改变碳分配以增加产量的方法,所述方法包括:表达UDP-葡萄糖差向异构酶,所述差向异构酶增加朝向膜生物合成的碳分配,由此减少可用于淀粉生物合成的碳的量。
11.根据权利要求10所述的方法,其中所述差向异构酶包含根据权利要求1所述的核苷酸序列。
12.一种植物,其在它的基因组中包含重组DNA构建体,所述构建体包含与至少一个调控元件可操作地连接的多核苷酸,其中所述多核苷酸包含根据权利要求1所述的核苷酸序列。
13.根据权利要求12所述的植物,其中与不包含所述重组DNA构建体的对照植物相比,所述植物表现出产量增加。
14.根据权利要求12所述的植物,其中与不包含所述重组DNA构建体的对照植物相比,所述植物表现出干旱耐受性。
15.根据权利要求12所述的植物,其中所述植物选自:玉米、大豆、向日葵、高粱、油菜、小麦、苜蓿、棉花、稻、大麦、粟、甘蔗、柳枝稷、烟草、马铃薯和糖用甜菜。
16.从根据权利要求12所述的植物生产的种子。
17.一种植物细胞,其在它的基因组中包含重组DNA构建体,所述构建体包含与至少一个调控元件可操作地连接的多核苷酸,其中所述多核苷酸包含根据权利要求1所述的核苷酸序列。
18.一种载体,其包含根据权利要求1所述的多核苷酸。
19.一种用于生产转基因植物的方法,所述方法包括:用重组DNA构建体转化植物细胞,所述构建体包含与至少一个调控元件可操作地连接的多核苷酸,其中所述多核苷酸包含根据权利要求1所述的核苷酸序列,和从所述转化的植物细胞再生转基因植物。
20.根据权利要求19所述的方法,其中所述植物选自:玉米、大豆、向日葵、高粱、油菜、小麦、苜蓿、棉花、稻、大麦、粟、甘蔗、柳枝稷、烟草、马铃薯和糖用甜菜。
21.一种增加植物的光合效率的方法,所述方法包括:表达根据权利要求1所述的重组多核苷酸,其中与不表达所述重组多核苷酸的对照植物相比,膜生物发生增加。
22.根据权利要求21所述的方法,其中所述植物选自:稻、小麦、玉米和大豆。
23.根据权利要求21所述的方法,其中所述增加的光合效率会导致增加的生长。
24.根据权利要求21所述的方法,其中所述植物表现出增加的籽粒产量。
25.一种寡核苷酸探针或引物,其源自SEQ ID NO: 18,且能够检测编码SEQ ID NO: 1的多肽的多核苷酸的表达。
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PCT/CN2011/000852 WO2011143933A1 (en) | 2010-05-17 | 2011-05-16 | Udp-glucose-4-epimerase useful for improving agronomic performance of plants |
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CN106636191A (zh) * | 2017-01-13 | 2017-05-10 | 中国科学院昆明植物研究所 | 拟南芥At‐UGE2基因及其过表达突变株和缺失突变株在调节植物性状中的应用 |
CN111172174A (zh) * | 2020-03-06 | 2020-05-19 | 沈阳农业大学 | OsUGE3基因在改善水稻性状中的应用 |
CN113755511A (zh) * | 2021-10-12 | 2021-12-07 | 河南大学三亚研究院 | 玉米Zm00001d029151基因在调控保卫细胞形态建成方面的应用 |
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CN109956996B (zh) * | 2017-12-14 | 2021-04-06 | 中国农业科学院作物科学研究所 | 一种谷子产量相关蛋白SiAMP1及其编码基因与应用 |
CN111172179B (zh) * | 2020-01-19 | 2020-09-08 | 武汉艾迪晶生物科技有限公司 | 泛素连接酶基因OsNLA2、蛋白及其在水稻选育中的应用 |
CN112322616B (zh) * | 2020-11-25 | 2022-04-05 | 上海市农业科学院 | 一种猴头菌来源的udp-葡萄糖-4-差向异构酶的表达和纯化方法 |
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CN106636191A (zh) * | 2017-01-13 | 2017-05-10 | 中国科学院昆明植物研究所 | 拟南芥At‐UGE2基因及其过表达突变株和缺失突变株在调节植物性状中的应用 |
CN111172174A (zh) * | 2020-03-06 | 2020-05-19 | 沈阳农业大学 | OsUGE3基因在改善水稻性状中的应用 |
CN113755511A (zh) * | 2021-10-12 | 2021-12-07 | 河南大学三亚研究院 | 玉米Zm00001d029151基因在调控保卫细胞形态建成方面的应用 |
CN113755511B (zh) * | 2021-10-12 | 2023-03-24 | 河南大学三亚研究院 | 玉米Zm00001d029151基因在调控保卫细胞形态建成方面的应用 |
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US20140059715A1 (en) | 2014-02-27 |
US9441217B2 (en) | 2016-09-13 |
BR112012029289A2 (pt) | 2017-05-02 |
WO2011143933A1 (en) | 2011-11-24 |
CN102250226B (zh) | 2013-06-12 |
CA2799050A1 (en) | 2011-11-24 |
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